Mycotaxon

- : «
- a ‘
; - - - .
‘ ‘ ; 2 - ee + r
a é = . -
> : 5 cad .
. 5 - — S ¢ : - % * fe -
x 5 . e "s 3 ¥ ‘
Ae + ind ray Ce ; :
= . > : j ( j
j 3 =o 3 ? , 2
" Whey 2 : . :
ee : : ; ‘ % - : : ‘ ‘ : +
3 a 2 = 4 . ¥
S “= bi F or x Bs —-
~ ey S - 0 4
; td : 5 j :
A . *. 3 é
5; 5 : a igs
; * ~ f So = “

ALBERT K. MANN LIBRARY
AT

COKNELL UNIVERSITY

CORNELL UNIVERSITY LIBRARY

INL VI

3 1924 074 216 494

«

)

42-

ie,

| MYCOTAZON

AN INTERNATIONAL JOURNAL FOR RESEARCH ON
TAXONOMY & NOMENCLATURE OF FUNGI, INCLUDING LICHENS

CONTENTS

A Catalogue of Types (Uredinales) in the National Mycological Herbartum (DAOM)
Roa ids AA os AR aac a feta les Una on el hackeye Gaunt eR aa J. A. Parmalee
Some Stemonitales (Myxomycetes) from the State of Tlaxcala, Mexico.
M. Rodriguez-Palma and A. Estrada-Torres
Studies on Mediterranean Hyphomycetes. Wl. Quadracaea mediterranea anam.-gen.

PIG ST ON ce oe end Su bie paneiance ela a cae D. Lunghini, F. Pinzari and L. Zucconi
Heterobasidiomycetes from Majorca & Cabrera (Balearic Islands)... ..... Peter Roberts
A New Species of Callistosporium......... Raymond M. Fatto and Alan H. Bessette

Scutellospora cerradensis: An Ornamented Species in the Gagasporaceae (Glomales)
from the Cerrado Region of Brazil.
Joyce Lance Spain and Jeanne Claessen De Miranda
Glomus brasilianum: An Ornamented Species in the Glomaceae.
Joyce Lance Spain and Jeanne Claessen De Miranda
Royoporus -A New Genus for Favolus spathulatus ... 0.2.0.0. es A.B. De
Ohytrormmyccice Laxonomy since 1900 ios ie a eta nets see Joyce K. Longcore
Coccoidella exocarpi sp. nov. on Exocarpos spp. from Australia.
Z. Q. Yuan, C. Mohammed and T. J. Wardiaw
New and Interesting Wood-Inhabiting Fungi (Basidiomycotina~Aphyllophorales)
APOE SEMA as er er deci 28 Kurt Hjortstam and Leif Ryvarden
A New Species of Bartalinia Tassi (Deuteromycotina, Coelomycetes).
Freda Anderson and Virginia Bianchinotti
Pinatubo oryzae gen. et sp. nov. and its Identity During Routine Tests of Rice Seed.
J. B. Manandhar and T. W. Mew
A Preliminary Checklist of Agaricales in Turkey ..... Eriugrul Sesli and Saffet Baydar
A New Species of Thermoascus with a Paecilomyces Anamorph and Other
Thermophilic Thermoascus Species from Taiwan.
Kuei-Yu Chen and Zuei-Ching Chen
Thielavia pingtungia sp. nov. a Thermophilic Ascomycete from Taiwan.
Kuei-Yu Chen and Zuei-Ching Chen
Hymenoscyphus seminis-alni, a New Species of the H. fructigenus - Complex.
Hans Otto Baral
Amoenodochium humicola anam. gen, et sp. nov., a New Sporodochial Hyphomycete
AOU OH La ean Fernando Pelaez and Rafael ¥. Castafieda Ruiz
Mycena sororia sp. nov., Close to M. rosea Gramberg (Basidiomycotina).
J. Perreau-Bertrand, M. C. Boisselier-Dubayle and J. Lambourdiere
[Contents continued inside front cover}
ISSN 0093-4666 MYXNAE 60: 1-524 Ree od

For subscription details, availability in microfilm and microfiche,
and availability of articles as tear sheets, see back cover.

Volume LX October-December 1996

[Contents continued from front cover]
Notes on Conidial Fungi. X. A New Species of Ceratosporella and Some New
CiiminhOns © onl hod Be es 2 R. F. Castafieda Ruiz, J. Guarro and J.Cano = 275
Some Airborne Conidial Fungi from Cuba.
R. F. Castafieda Ruiz, D. E. Fabré, M. P. Parra, M. Pérez and J.Guarro 283
Egyptian Uredinales. I. Rusts on Wild Plants from the Nile Delta.
Zakaria A. Baka and Halvor B. Gjerum 291
The Genus Pyrofomes in Argentina and the Curltural Features of P. lateritius

(Aphyllophorales).
Jorge E. Wright, Marcela F. Bolontrade and Alejandra T. Fazio 305
Taxonomy of Fomitopsis rubidus comb. nov.......---- Anjali Roy and A. B. De 317

Strigulaocculta, a New Saxicolous Lichen from New Zealand.
P. M. McCarthy and W. M. Malcolm 323
Xylaria Species from Papua New Guinea: Cultural and Anamorphic Studies.
Katleen Van der Gucht 327
A New Species of Hygrocybe from India.
T. K. Abraham, K. B. Vrinda and C. K. Pradeep 361
NATS Truffle and Truffle-Like Fungi 5: Tuber lyonii (=T. texense), with a Key to the
Spiny-Spored Tuber Species Groups.
James E. Trappe, Ari M. Jumpponen and Efrén Cazares 365
Hyphomycetes on Alchorneatriplinervia (Spreng.)Miill. Arg. Leaf Litter from the
Ecological Reserve Juréia-Itatins, State of Sao Paulo, Brazil.
Rosely Ana Piccolo Grandi and Derlene Silva Attili 373
Bridgeoporus, a New Genus to Accommodate Oxyporus nobilissimus (Basidiomycotina,
Polyporaceae).
Harold H. Burdsall, Jr., Thomas J. Volk and Joseph F. Ammirati, Jr. 387
A Check List of Higher Marine Fungi on Wood from Danish Coasts.
J. Koch and K. R. L. Petersen 397
Lichenicolous Fungi from the Western Pyrenees, France and Spain.

II. More Deuteromycetes........-......- Javier Etayo and Paul Diederich 415
A New Species of Botryobasidium (Corticiaceae, Aphyllophorales) Inhabiting the
Interiors of Brown Rotted Logs of Ponderosa Pine.......... Michael J. Larsen
Alan E. Harvey, Robert Powers and Martin F. Jurgensen 429
Planistromelaceae, a New Family in the Dothideales............. Margaret E. Barr 433

A New Species of Alysidiopsis from Mexico.
Angel Mercado Sierra, Maria José Figueras, Julio Mena Portales 443
New and Interesting Helicoon Species from Spain.
Samir K. Abdullah, J. Guarro and M. J. Figueras 449
Hyphomycetes from Streams: New Taxa and New Combination.
Ludmila Marvanova and Enrique Descals 455
ALLRUS: A System for Standard Description, Identification and Classification
of Russulaceae. V. Robert and B. Buyck 471
Helicoon septatissimum sp. nov., a New Species from Tierra del Fuego.
A. M. Godeas and A. M. Arambarri 481
A New Species of Leptodontidium from Quercus acutissima.
A. Tsuneda, N. Maekawa and R. S. Currah 485

Rica TEC IOWE ics a oo a DO ho. 8 ib Bs ps Few Oe ee a ee els D. S. Hibbett 491
NOTICE: Transfer of Professor E. J. H. Corner's Mycologial Material ........-..---- 497
Pasir WIEN fds ak. A atin Bales ic ok 6x de BAe eS ee Fe oe ie eee 498
Index to Fungous and Lichen Taxa... . 2... 6. eee eee eee eee tee eee eens 501
PP OULW EE co 6 hick EE Oh Se Be ee bes DEAR licks ot Mieg oh > Oa eae eee 524
Publications Date, Vouutne SO i462 os ecb ot oes ww eles we BieLae + eee 524

© Mycotaxon, Ltd., 1996

MYCOTAXON

AN INTERNATIONAL JOURNAL OF RESEARCH ON
TAXONOMY & NOMENCLATURE OF FUNGI, INCLUDING LICHENS

VOLUME LX, 1996

COMPLETE IN ONE VOLUME
CONSISTING OF iv + 524 PAGES INCLUDING FIGURES

EDITOR-IN-CHIEF

Jean Boise Cargill
Harvard University Herbaria, 22 Divinity Ave.
Cambridge, MA 02138-2094, USA

Pavel Lizon
Assistant to the Editor-in-Chief
Plant Pathology Herbarium
Cornell University, Ithaca, NY 14853

ASSOCIATE EDITORS

David S. Hibbett
Book Review Editor
Harvard University Herbaria, 22 Divinity Ave.
Cambridge, MA 02138-2094, USA

Grégoire L. Hennebert
French Language Editor
32 Rue de I'Elevage
B-1340 Ottignies -LLN, Belgium

Karen Gettelman
Index Editor
Indiana University
Bloomington, IN, USA

EDITORIAL ADVISORY BOARD

JAMES W. KIMBROUGH, Gainesville, Florida (1992-97, Chm.)
AMY Y. ROSSMAN, Beltsville, Maryland (1990-96)
GREGOIRE L. HENNEBERT, Louvain-la-Neuve, Belgium (1990-96)
JACK D. ROGERS, Pullman, Washington (1993-98)

OVE ERIKSSON, Umea, Sweden (1994-99)

RONALD H. PETERSEN, Knoxville, Tennessee (1995-2000)

Published by
MY COTAXON, LTD.P.O.BOX 264
ITHACA, NY 14851-0264, USA

Printed in the United States of America

© Mycotaxon, Ltd., 1996

iii

Table of Contents, Volume Sixty
October-December 1996

A Catalogue of Types (Uredinales) in the National Mycological Herbarium (DAOM)
AWA ANada Le ec ie ee ee ee ke J. A. Parmalee 1
Some Stemonitales (Myxomycetes) from the State of Tlaxcala, Mexico.
M. Rodriguez-Palma and A. Estrada-Torres 79
Studies on Mediterranean Hyphomycetes. III. Quadracaeamediterranea anam.-gen.
ANCES HOV GT iene ake et ce ons D. Lunghini, F. Pinzari and L. Zucconi 103
Heterobasidiomycetes from Majorca & Cabrera (Balearic Islands). . .Peter Roberts 111
A New Species of Callistosporium. . Raymond M. Fatto and Alan E. Bessette 125
Scutellospora cerradensis: An Ornamented Species in the Gagasporaceae (Glomales)
from the Cerrado Region of Brazil.
Joyce Lance Spain and Jeanne Claessen De Miranda 129
Glomus brasilianum: An Ornamented Species in the Glomaceae.
Joyce Lance Spain and Jeanne Claessen De Miranda 137
Royoporus —A New Genus for Favolus spathulatus ............4.. A. B. De 143
Chytridiomycete Taxonomy Since 1960................ Joyce E. Longcore 149
Coccoidellaexocarpi sp. nov. on Exocarpos spp. from Australia.
Z. Q. Yuan, C. Mohammed and T. J. Wardlaw 175
New and Interesting Wood-Inhabiting Fungi (Basidiomycotina—A phyllophorales)
POMELCIODIA TT rare or Kurt Hjortstam and Leif Ryvarden 181
A New Species of Bartalinia Tassi (Deuteromycotina, Coelomycetes).
Freda Anderson and Virginia Bianchinotti 191
Pinatubo oryzae gen. et sp. nov. and its Identity During Routine Tests of Rice Seed.
J. B. Manandhar and T. W. Mew 201
A Preliminary Checklist of A garicales in Turkey.
Ertugrul Sesli and Saffet Baydar 213
A New Species of Thermoascus with a Paecilomyces Anamorph and Other
Thermophilic Thermoascus Species from Taiwan.
Kuei-Yu Chen and Zuei-Ching Chen 225
Thielaviapingtungia sp. nov. a Thermophilic Ascomycete from Taiwan.
Kuei-Yu Chen and Zuei-Ching Chen 241
Hymenoscyphus seminis-alni, a New Species of the H. fructigenus - Complex.
Hans Otto Baral 249
Amoenodochium humicola anam. gen. et sp. nov., a New Sporodochial Hyphomycete
from Indian Soil. ..... Fernando Pelaez and Rafael F. Castafieda Ruiz 257
Mycenasororia sp. nov., Close to M. rosea Gramberg (Basidiomycotina).
J. Perreau-Bertrand, M. C. Boisselier-Dubayle and J. Lambourdiere 263
Notes on Conidial Fungi. X. A New Species of Ceratosporella and Some New
Combinations; -) 052% R. F. Castafieda Ruiz, J. Guarro and J. Cano 275
Some Airbome Conidial Fungi from Cuba. R. F. Castafieda Ruiz, D. E. Fabré
M. P. Parra, M. Pérez and J. Guarro 283
Egyptian Uredinales. I. Rusts on Wild Plants from the Nile Delta.
Zakaria A. Baka and Halvor B. Gjzrum 291
The Genus Pyrofomes in Argentina and the Curltural Features of P. lateritius
(Aphyllophorales).

Jorge E. Wright, Marcela F. Bolontrade and Alejandra T. Fazio 305
Taxonomy of Fomitopsis rubidus comb. nov....... Anjali Roy and A. B. De 317
Strigulaocculta, a New Saxicolous Lichen from New Zealand.

P. M. McCarthy and W. M. Malcolm 323
Xylaria Species from Papua New Guinea: Cultural and Anamorphic Studies.
Katleen Van der Gucht 327
A New Species of Hygrocybe from India.
T. K. Abraham, K. B. Vrinda and C. K. Pradeep 361

iv
NATS Traffle and Truffle-Like Fungi 5: Tuber lyonii (=T. texense), with a Key to the
Spiny-Spored Tuber Species Groups.
James E. Trappe, Ari M. Jumpponen and Efrén Cazares 365
Hyphomycetes on Alchorneatriplinervia (Spreng.)Miill. Arg. Leaf Litter from the
Ecological Reserve Juréia-Itatins, State of Sdo Paulo, Brazil.
Rosely Ana Piccolo Grandi and Derlene Silva Attili 373
Bridgeoporus, a New Genus to Accommodate Oxyporus nobilissimus
(Basidiomycotina, Polyporaceae)..........--: Harold H. Burdsall, Jr.
Thomas J. Volk and Joseph F. Ammirati, Jr. 387
A Check List of Higher Marine Fungi on Wood from Danish Coasts.
J. Koch and K. R. L. Petersen 397
Lichenicolous Fungi from the Western Pyrenees, France and Spain.
II. More Deuteromycetes...........-. Javier Etayo and Paul Diederich 415
A New Species of Botryobasidium (Corticiaceae, Aphyllophorales) Inhabiting the
Interiors of Brown Rotted Logs of Ponderosa Pine ...... Michael J. Larsen
Alan E. Harvey, Robert Powers and Martin F. Jurgensen 429
Planistromelaceae, a New Family in the Dothideales......... Margaret E. Barr 433
A New Species of Alysidiopsis from Mexico.
Angel Mercado Sierra, Maria José Figueras, Julio Mena Portales 443
New and Interesting Helicoon Species from Spain.
Samir K. Abdullah, J. Guarro and M. J. Figueras 449
Hyphomycetes from Streams: New Taxa and New Combination.
Ludmila Marvanova and Enrique Descals 455
ALLRUS: A System for Standard Description, Identification and Classification
of Russulaceae. V. Robert and B. Buyck 471
Helicoon septatissimum sp. nov., a New Species from Tierra del Fuego.
A. M. Godeas and A. M. Arambarri 481
A New Species of Leptodontidium from Quercus acutissima.
A. Tsuneda, N. Maekawa and R. S. Currah 2

Book Reviews wise as ibek weet abe s ms Noaeitie Jats suise ietclier olabten aca a D. S. Hibbett 491
NOTICE: Transfer of Professor E. J. H. Corner's Mycologial Material.......... 497
‘Authonindex 3.000505 6 osteo osc neIED cules BP tae aas: 4) lek Wan sacte eailice ela 498
Index to Fungous and Lichen Taxa ... 2.2.2.6 - eee cree eee cee cee e en ees 501
REVIEWETS 650. 5 He alee sats os 4 meses no ale ale tae ok se ea ar heels (aac aa 524

Publications Date: Voliine 59/35) 298 eos ap nu are ee ieaite ie oo cars as eal ear ol et arieayy aetlal ae 524

MYCOTAXON

Si NN SO Ns IRD tl RA ARTS STAR EN it eS tI a
Volume LX, pp: 1-77 October-December 1996

A CATALOGUE OF TYPES (UREDINALES) IN THE
NATIONAL MYCOLOGICAL HERBARIUM (DAOM)
OTTAWA, CANADA

J.A. PARMELEE

Centre for Land and Biological Resources Research
C.E.F. Ottawa, Ontario, Canada KIA OC6

¢ ABSTRACT
A search for type specimens of rusts in Canada’s National Mycological
Herbarium found over 500 amongst the 50,000 specimens of rusts. These are
recorded in tabular format indicating hosts (and families), DAOM accession
number, exsiccati name and number if applicable, collector and number if
available and the specified type.

INTRODUCTION

At a symposium on taxonomy of rust fungi at the 4th
International Mycological Congress in Germany in 1990, it was proposed
that a record of nomenclatural type specimens held in various herbaria
would prove a useful aid to workers in this field. Herein is a record of
all such types in the National Mycological Herbarium (DAOM). Fungus
collections began in DAOM mainly with the appointment of the first
Dominion Botanist for Canada in 1909, but holdings today date even
earlier with the purchase and exchange of specimens from Asia, Europe
and the United States. Authors of some of our larger exsiccati include:
Ellis and Everhart (Fungi Columbiani 1-5100 and North American Fungi
1-2900), Rabenhorst (Fungi Europaei 1-4100), Petrak (Mycotheca
Generalis 101-2100), Bartholomew (North American Uredinales 1-3500),
Sydow (Uredineen 1001-2650), Thuemen (Mycotheca Universalis 1-2300).

Z,

One smaller exsiccati obtained in the 1950’s dates from 1810 by Mougeot.

Some of our exchange material includes specimens from Holway
(Mexico, Central and South America), Arthur (North America),
Cummins (North America), N. Hiratsuka (Japan) and H.S. Jackson
(South America). At this writing the National Mycological Herbarium
contains over 300,000 specimens of all major groups of fungi and I
estimate that 50,000 are rusts. In this review I found about 1% to be
types - 629 by count.

As outlined in the International Code of Botanical Nomenclature
(ICBN) W. Greuter et al. (1994), nomenclature in the fungi dates from
1753 but names of rusts adopted by Persoon (1801) are sanctioned i.e.
treated as if conserved against earlier homonyms and competing
synonyms. Nomenclatural types as delimited in ICBN Art. 9 include:

HOLOTYPE - the one specimen or illustration used by the author, or
designated by the author as the nomenclatural type.

ISOTYPE - any duplicate of the holotype.

SYNTYPE - any one of two or more specimens cited in the
protologue when no holotype was designated.

PARATYPE -_ a specimen cited in the protologue that is neither
holotype nor an isotype, nor one of the syntypes.

LECTOTYPE - a specimen or illustration designated as_ the
nomenclatural type when no holotype was designated.

NEOTYPE - a specimen or illustration selected to serve as
nomenclature type as long as all of the material on
which the name of the taxon was based is missing.

TOPOTYPE - _alater collection from the type locality (ex. ICBN 1981).

Types reported here have been transferred to specially marked
standard herbarium sheets with bold cross-hatching about 2-3 cm. high
completely across the bottom of both sides. These sheets are restricted
for a type specimen and any notes, photographs, drawings or slides from
same plus the original description.

Generic names of rusts included are based on Cummins and
Hiratsuka (1983). The specific and subspecific taxa are as they appear on

3

the labels, but if now known to have been reduced to synonomy, the
recognized epithet is included in parentheses (= .....). Doubtless other
taxa may prove to be synonyms also. Basionyms are likewise included in
parentheses but without the equals symbol. The genera and species are
listed alphabetically but in those genera having numerous species, like
Puccinia and Uromyces, the species are listed alphabetically within host
families. For example Puccinia shiraiana is the fourth species listed
under the first alphabetical family Acanthaceae, P. arabicola is the
second species under Brassicaceae and P. aegopogonis is the first of 35
species under Poaceae. Taxa of Puccinia on Asteraceae are numerous
(175) and host tribes in this family are also sequenced
(Asteraceae/Anthemideae, Asteraceae/Astereae, Asteraceae/Cardueae....)
with rust taxa alphabetically under each tribe. Employment of hosts in
this manner reflects arrangement in DAOM. Included under the
remarks column are: the DAOM number, the collector’s name and
number or an exsiccati name and number, occasionally a related note and
a designation of type in bold face. Delimited types are given when clearly
obvious otherwise as indicated on the specimen label e.g. type material,
part of the type or simply type. Herbaria are designated as in the eighth
edition of Index Herbariorum by P.K. Holmgren et al. (1990) and the
private collection of N. Hiratsuka is abbreviated as HH following the
treatment in N. Hiratsuka et al. (1992). Authors of rust taxa cited
regularly throughout are abbreviated: J.C. Arthur (Arth.), G.B.
Cummins (Cumm.), P. Dietel (Diet.), Nahoide Hiratsuka (Hirats. f.),
E.W.D. Holway (Holw.), H.S. Jackson (Jacks.); otherwise names are
abbreviated according to Arthur and Cummins (1962) or spelled in full.

ACKNOWLEDGEMENTS
For their reviews of the manuscript, I am grateful to Dr. Y.
Hiratsuka, Northern Forestry Centre, Edmonton, Alberta and Dr. G.B.
Cummins, Tucson, Arizona. In-house reviews were by Drs. J. Ginns and
S.A. Redhead and I also acknowledge the aid by staff of the National
Mycological Herbarium, Ottawa in preparing camera ready copy for
publication.

REFERENCES

Arthur J.C. and G.B. Cummins (1962). Manual of the rusts of United
States and Canada. Hafner, New York. 438 pp.

Cummins G.B. and Y. Hiratsuka (1983). Ilustrated genera of rust fungi -
Rev. ed. American Phytopathological Society, St. Paul, Minn.
152 pp.

Greuter W., F.R. Barrie, H.M. Burdet, W.G. Chaloner, V. Demoulin,
D.L. Hawksworth, P.M. Jorgensen, D.H. Nicolson, P.C. Silva,
P. Trehane, J. McNeil (1988). International Code of Botanical
Nomenclature. (Tokyo Code). Koeltz Scientific Books.
KGnigstein, Germany. 389 pp.

Hiratsuka N., S. Sato, K. Katsuya, M. Kakishima, Y. Hiratsuka, S.
Kaneko, Y. Ono, T. Sato, Y. Harada, T. Hiratsuka and K.
Nakayama (1992). The rust flora of Japan. Tsukuba Shippankai,
Tsukuba, Japan. 1205 pp. + index 159 pp.

Holmgren P.K., N.H. Holmgren and L.C. Barnett (1990). Index
Herbariorum 8th ed. Regnum Veg. 120:1-693.

Persoon C.H. (1801). Synopsis methodica fungorum. Gottingen. 706 pp.

[edAy
Jo ued] IWI X9 7877 UoIysieg ‘TyE61 WOVG

[edAjosy] DLL X9 OL9T “sxoRL ‘86807 WOVAd
[adAjos]] UNd X9 €6€E"7 WOVA

[adAj0s]] €pZ1 “MIOH ‘Z109L WOVG

[ad4josq] snowkag ‘eweqely ‘gz7~9 WOVG

UNd 01 sdAjos: [adAjopoxy] BIAeg ‘716€ WOVA

SWAVNGAa

‘ds vai0osoig
"J sudosajNy BAT
[[J9py PIOYNINUIZ eWOsOI[s]

‘ds ensesnbnya

‘Od ("7) susssany eryso0g

(winsosn
winuiojedngq =) ‘suiqoy
2p Bury (“T) eulssnye euntiesy

LSOH

‘UIUIND SUdUO0D]
"syour oval

‘wUuInd WNuUTOUI0Y

“MIOH 27 ‘syorg sensesnbnyo

*pAg oeIysis10q

Qplaeg seunesose

WNIdIodav

IVINONIG

WOVG NI (SATVNIGAYN) SNOLLOATIOO AdAL

[adAjos]] Zp “MIOH ‘1009L WOVA

[adAjosq] J2nf "CO *9L09L WOVG
[edAjosq] 91TZ1 “10H ‘7L09L WOVG

[adAjuAg] STIS “MIOH ‘ST8801 WOVd

[adAjosq] ¢9¢ “MIOH ‘09€8L NOVA
[adAjosq] gOET “MICH “¥909L WOVG
[adAj0s]] ge “MIOH “€909L NOVA
[adAjosq] 6€€T “MIOH “6S09L WOVG
[adAjosq] €9 “MIOH “8S09L WOVG

[adAjos]] 99¢€1 “MIOH ‘SS09L WOVA
[adAjosq] e6pZ “MIOH “pS09L WOVG

[adAy, Jo wed] g¢z7z 94D “Eo8er WOVA

(‘3aN 29 “321d
aellaweonse “y =) ‘ds eliawaons[Ty

“J eouedsies visdeyy

avooeyUeIeUYy

[eH 2 “Uey] sisuodeyex eipuey

SII, sisuayinb ewosysoj1990d
“MEPL PI[OJIPULIS eIySoIEg

‘d 3 “Y Wmyeulouns wnuerjos
“Sse eIpoytsoddo evipuejseN
[Yq umNjjayoqnd umnuejos
“quNY], BIOJSeIIIN eIADNS

Joyeg eueisiddood eipiWw

‘ds wnonsn3i]

“MJOH 2 “SyORL Wnsosqoua}

Jeng wntorund

"syorr ovjeid

‘yy wryusnieatnd

“MOH
w ‘sores aewoysosios0d

“MJOH 2 “syour oerysosod

“MJOH 2 “SyoRL sisuopnded

“MJOH 2 “SyoeL ovspue}ou

“MOH 2 “syoes e[Npunus
“MOH 2 “syoer WNUTUTU
“MOH “SYOeL sey[IW
‘WwUIND BjOOLNsNsI]

°p,jUOd MIMNIPIEy

[adAjosq] 18 910 2 “ZION ‘LTI9L WOVG

[sed43osq] WOV Ut €00I “19S
“"IEY “OIA! “IS2A = O9TS “MIOH ‘618801 WOVAd

[adAjosq] cozs “MIOH ‘818801 WOVAG
[adAyosy] 7967 ystieg “€77691 WOVA

(€861) ‘SIBIN *“A % “wWuINnD spy erluosouWAD
Joj snuas juoweoedel ‘WOVQ Ul jou adAT

[adAjosq] 9401 “MIOH ‘€119L WOVA

“J stiqnjoa vayessoddizy

"MAH Seploliessuld o1dauas

“SAH 9suerenszed wintojyedng

‘Od Isse[snop o1saueg

“JT sijeyuapioc0 snqny

‘ds u0}01D

eATO
wp [EZIOUAA Bea}eID0dd1y

VZIHXYOAALO

“yy rAemjoy

(‘9 erfaia1q)
“yuy (ypy) Woyedna

“Ey snomtojiyes

SNWOUGOAV A

(‘d eruosouuwlAy)

“Sve A 2 “wun
(‘Yq Ul aMOH) euRTyDOd

SHOAWOMNALAV

“MJOH 2 ‘Syoes Benue

VIMNH.Lav

S ‘IW ‘OLYL “UNd ‘HOIW ‘Idd
0} sadAjost ‘["1ea Jo adAjojoH] ZrpSp WOVA

S ‘O ‘IWI ‘OLUL “ANd “HOIW “HA ‘Idd
0} sodAjost ‘["1ea Jo adAjojoH] p8pSp WOVA

[edAje1eg]
6€8 IIRNON ‘preppy UNd X9 E9Se7 WOVA

[adAjosq] 6ST “MIOH ‘1S19L WOVG

[edAjos]] CTET “MIOH ‘TPI9L WOVG

[adAjos]q] [I pu eS “L ‘PO8I61 WOVG

JapaQ Wndipurjus0I3 wWNnpsy]

‘YNNY Wnsojnpurys winpsyT

XI]

“OC Bojorra[ppng eluexAY

ovoorluoUsig

‘quny], Beyjaquin snuseoryq

otAes
IOIpUP[UZOIS “IA

aJIAvs IsO[Npueyls “Iva

Areg
3p (MYO 2 “QIY) IPel

("1 WINIpIooy)
“UY CAG 29 “1D eUloIy!

VXAWOSAYHO

(‘wi elosdosAry)
"pAg Cypry) oerueyu

SNTOADOSAYHO
uosyoer IAeMOY

WNiTaLOwdo

o1eg *s
FL ‘eunysiyey lwsaxje

VaOSdOdOVAHO

[addy Jo used] p90 “MIOH ‘€179Z WOVG

[adAjuAg] 1OSE “MOH ‘€Z78801 WOVa
[j"Josop jou eI]AI] GPST “SydeL ‘SO7TIL WOVA

[ed Ajos]]
1681 “uel ‘unsylese’T ‘7079 WOVd

[adAjosq] 127 9M ‘€619L WOVA

68LL dHAVC Ut aedAjofoxy
‘[edA4je18qg] 6$LL dJAVG = O1S9E WOVG

WNd 0} osye odAost
‘[1ea Jo adAj0sq] OSP6 dIAVG = 00S9¢ WOVG

‘ds eluosog

Aely “y ejeydacosoeyds eutsaqioA

*YOOH JO[OIIq Uo|Ne.OUspYy

*ssec e][oyond stxouAy

Auieg Wuewyasua essig

Kely “Vy ("J) eioyiun sasauojy

"WS “a'r Wintpoytased winturs0e A,

‘iy oeIU0seq

“MOH
Ul “M[OH 2 ‘JoIq sdooue

"syour stuo[nesouape

WhldOdSOuTtoo

“"ysose’y] siprxoud3

VuOSdOSAUHO

“SOUL ILIIOM

JOIIIZ sisououw

JOTIVZ WIDVA “IBA

*p,juos exAwosd143)

10

[edAjodoy)] 60SSL VASN x9 88€9L WOVG

[adAjosy] ‘u's “MOH ‘OP8801 WOVa
[ad4juAg] ZpSe “MIOH ‘pEs9L WOVA

[eddy jo wed}
WOVG U 9001 “3919S “JeY “OIA “"IS8A

[adAy, Jo ued] O18 O2Y4D “prscy WOVA

[adAjosq] Eye “MIOH ‘SE880I 2 6LZ9L WOVA
[edAjosT] WOV U! €S6 “ON “AUN “oA< S'WANYL,

OWLW x2
[sadAjosq] €¢ [19401 “L900 PF 9PEET WOVG

[adAjosy] 907 SuenyZ ‘peels WOVd

[addy Jo ued] gps osyD ‘6sscp WOVA

ysing winaine soqry

"M@H seplowpuersy eras,

“ds vulsaqioA

“YOOH (‘DG) Saplorjayorn erAerg
“J erOJIpioo eIqny

"YOOH 2 “‘Wueg Jojoosip winger]

"PITLAA Snuvautjoies sndojueydalq

Souler SI[IXo[J SNUIg

WH 29 Ing
(‘qxoy) sleyixe seipuodsols0ya

“ds snajoa

wn 2 [oped
‘“-93poH{ o[BIUSPI9I0

WNILLYVNOWO

“MJOH 2 “IC VeIOINIIA

"JQIC] DBUISOQIOA

"ULIY deIAa}S
"WUIND ejOoIgnI

“MJOH Ul “MOH
2» ‘JIC WMeseydesed

‘wonyy sipodojueydays

"UNWIND H]JaMolo

suenyZ
2 Sues, sueIpuodsoss0yo

‘wun wWnueOSYyS

*p,juos umntodsoajod

11

[adAjosq] Z78¢ “M1OH ‘ppr9L WOVA

[jetzajyeur ad] COPT “MIOH ‘67P9L WOVA

[adAje1eg] 1681 “Idv
OLOT esual|iseig “qiaH ‘eI “| ‘97P9L WOVA

[lelzayew adAy] peZI “MIOH ‘8EP9L WOVAG

[adAj0s]]
U0}00M ‘O'F “8061 “8NV 6 ‘61P9L WOVG

[adAjos}] WOV UI c6cT ‘doing I3unz ‘“-qey

Bolyodyes

"TH ‘I$ stusoo1suo] enaywinisy,

"yeg esojuoWio) enselnbnyD

‘ds esuy

opped
(U0}00M) ediesO;EUIDeY eIUOYRY

ysing winoine saqry

(‘y uinjAydopug) “uuazy 2
‘Ing (‘syoer) IAeMOoy

VITALaIda

“MOH
wz? “syoers senazuniy)

[6681

PST:8E eISIMpSH ‘91d
99S] “13Iq senseimnbnyo

VaOSdOWACIG

“MIOH 2 ‘syorg eqiodns

VINISIGHOId

(‘mM stxAdoig) ‘yy
(‘yyy) eurIuO}00M

VTTHISNINWNO

JOYOSL] B[OOIQl

*p,juod WINTIeEUOID

L2

[edAje1eg] SIAeq ‘f'f
“OSIM “TT PUIAA “TLPOL puke 878L WOVG

[adAjo;0H]
SIQUUOD 2 61S se]9WIed ‘Z717SL WOVA

[edAjuds} HA X29 ~Z796L1 WOVAd

[[erojeu
adAy, xa] aseurep yoasut Aq 100d uoumtdeds

‘QESOI UNd Xe (AjuO sajou) geeeelT WOVA

Bireqioy] SsnoJowNu 0}
sodAjost ‘[adAjojopy] 9¢ZE BLAeS ‘COP9’ WOVA

[adAjojoH]] &7879 JOpleD “EILSS WOVA

[ietsajyeur dA] pos “MIOH “8Ep9L WOVA

“J stunwiwos sniodiune

yousopy] SI]BUOZIJOY snodiuns

“J euRIUIZIIA sniodiune

*Z1eg wnJojndoos sniodiune

*XYOIJAL CULIUTTOIeD BIUOIARID

winuldje “q se
"WET WINTOsIpITeseue umnigopidg

Aqsny eyeoiealp sioqiog

Woy HSIAep

sojsUuLIeg WSJOUUOD

gjieq wnueipnuieq

wley 1ayeq
WAIDNVAOdSONWAD

sofoulleg
W IIAeS SIsoJ-snoe]

ayaeg winueyseye

WNTIAHdAOGNG

"MJOH 2 “syoee eI[OuUs}

VoHLAGH

13

[edAjodoy]
6006 OSIeg ‘7978 UID “QIeH = LpS9L WOVa
[edAy
jo ied] OLOTT UNd = (AlUo apis) TpEEeEel WOVG
[adAjodoj]

ILZS ISH “L066 UNd = 996L91 WOVA

[addy Jo ued] WOV UI 9 ‘sxq “Joxq 13uny
“pg xa ‘(ATUO apts) ZPEEEl WOVA

[dss Jo adAjojo Fy]
ByNsHyeNH “AX WY COCE sojouleg “870971 WOVd
[adAy Jo wed] younz ‘aynyosysoyy
"YooL “wus “jog ul odAT, x2 996811 WOVG
[edAjosy] WOV U! 6LpT 13unq ‘Wy “N “II

uounseds
UI SNOIAQO 3Nq PaqlJOsap 10U ITS} :3)}0U
[edAjos]] WOVA UI [LZ I3unq ‘wy *N “II

[ad], Jo ued) 61ITI UNd
xo ‘(AjUO apl|s) OPEEET WOVG

‘yoeds
(que) stUdeIeyJ00U sIIedAdaeWIeEYD

‘31eg wnsojndoos sniediune

*YOOH Sijejuapiss0 sniodiung

sIsuaUTYS sniodiung

""T stunumwos sniodiung

“PIAA eueu sniodiune

‘dS CD saprodwp stuedAsaeueyD

“d'S'd (CD saprokwp stuedAdoewreys

‘qpAy sisuoyejn eurqes

(‘U Opeln)
"UV (‘[21,L) ssuajeyIOou

"yy Wuosjau

‘yy wntueissouysey

(epewiex xo
oqeAIP WNdHeIse “D =)
*‘pAS winuvoesey

do[oueg
sisuazioqye “dss

3307 lWuewnees

UJOY WNUII}eI

STA Hst|T9

way wninp

*p.juod winisue10dsouuhs

14

(‘ZISI ‘687 ‘ou sewmesojdAIZ
sading ‘JapISON 2 JOosNOP] SI Jsatjiee Inq
WOVC UI S,08g81 Wor sususeds uvadoing Aueuw]

[adAy jo wed] ¢91 HH X? pr867 WOVA

[edAjosq] 1691 ‘Pesn “PAS = L9S9L WOVG

[adAy, x9] projsuey ‘L0€761 WOVAG

[adAy jo ued] 9¢€01 HH X? “LIL7TE WOVA

[edAjosq] WOVC UF ZS “AUN, “OoAW “WaNnyL

sijigelj stiajdoAIq se
"yulog (7) sipsey elsajdoyskZ5

‘\slIyD UNJasIwenbs winizeldiq

"Jy slisus-tfides wnyueipy

‘Iq snsoiseds snyuekxO

“WIXR, SNyJoUTjOEd snqny

"Wg sNpIsi snqny

*$1aq UpodAjod q “IBA SLIvsUl]
Opeip) se ‘use upodAjod

‘J “syestH Wzeydip

(‘8 opain) “PAs (Dd)
SIUIOA-I]][1ded-URIpPE

VuOSdOTVAH

‘wuins 1mURAXO

VIG TINGH

"J ‘S}eIIGY Stu90Z0

(‘| wniprusesyd)
‘WIOy (‘Won ]) eWISsIsUO]

VuOdSVNVH

[edAT] SOE ‘syoRL ‘9OL9L puke SOL9L WOVA

[ed you nq WOVG UI

(OI8T) UONDaIO9 ysarpIea SI gIZ8S WOVG
ul 06 “IdAID sading ‘Jepsen 2 J093NoP]

[ds -J Jo adAjos]] £796 dIAVC X2 LOP8S WOVA
[adAy, Jo wed] O¢Z C2Y4D ‘pegep WOVA

[edAjosq] O18 “MOH = ZI99L WOVG

[sadAjereg] 07 “MICH = 9099
‘CTL “MMIOH = S099L ‘VELO “MICH = £099L WOVA

[edAjosq] €96 “MOH = €099L WOVG

[adAjojoH]] Se9881 WOVG

Jodig (‘YOOH) ele8nas eliedsig

“Jy wnonsepes wnury

yeleg euellgjnoos xyes

‘ds soywinapy

"MH Sisusj030q snqny

‘dds snqny
‘ds snqny

J “YOOH 2 “usg
(UIY 27 I4311A\) eURJUOW eILIOdsouWAD

"syour oerseosid
(Od ‘1 Open
se) “Aa] ((quosryq) tut

JaqVZ oe3ns}
‘ds ‘J ‘wonyy, vayida

"WUIND SsIptnaye

VaOSdWV TAW

"MOH 2 “SHEL eyjausy

"MOH 2 “syoee WAeMoy
"MOH 2 “Syoer viejo

VISNIVW

“UBJIN 2 ‘UBASeg IIeYyoeUe]

VTIOUNHANY

16

[edAyosq] (ZE6T) Ine Aq paso
se Use ‘q eURLIOZALIY efjososdwejayy

qns ‘WOVG UI pE9T “PAIN “PAS = 78L9L WOVG

[edAy,
jo ued] 81] HH X? ‘J “SIeIH °66L67 WOVG

[adAjodoy] towey ‘1849. WOVd

[adAjyodoy] tpuejedoss
Opesn qns ossT “pein ‘PAS = 79L9L WOVG
[adAy, Jo ued] [11 HH X® 76L67 WOVA

[addy jo ued] ¢, HH X® $167 WOVA

OZSZ “U,1109 epIyon & Woy Inq “paquiosap
jou Ba) :aI0N “fedA Jo ued] gp/67 WOVG

[adAjosq] susy90 ‘8871 ATH = 969191 WOVG

uinsoynurds

winipidsy qns syonj ‘d’H
(TEA) Buvisnyes siiadoAIg

"yey aiqewe wnuyse|g

“J o1e3[NA winipodAjog
“UNJOH

(7) ajeuolnuajdes winiusldsy

"WS “¢ loUNjIO} WNIWIOVIAD

eouodel
"rea eotuoddiu sisdoyuesids
se ‘yey wnoiuoddiu wnuyseg

‘qunyy, wnsioul wniusldsy

"1SI9O
(‘1pugq 27 ‘ddaog) eurdje snsejompon

(“¥ eyjorosdurepay))
“Use (‘UBeJ) PULLIOZOIIY

‘J ‘syeqlp] eULIOWey
J
“SyeIIE] 2p lowly sisusozaf

‘use (USeP) HYyotInay

emnwio—
e J sweny eueryney

“J "SyeITE] EjOoUYDe|q

(‘e BISopIAD “J “SIeIIH
(\jne.J) Bstoul-uaydse

VNISW TIN

U0sI3}0g W SUdTYIO eQIe

VIWADANOWIN

17

[eddy Jo ued] [¢9¢ HANL X? €618L1 WOVG

[ad4jos]] Wova wu
"y wniwserydiuy se pp] ‘sxq ‘oxq “4 ‘*pAs

[edAy, Jo y1ed] eSeurysox ‘66,9, WOVA

[adAjos]q] 90ue1,7 ‘sas0A jenUS. WOY
“818 “Wa “SAW “PAS = 76L9L % 16L9L WOVA

[adAy,
jo ued] OyT HH x2 eseurqol ‘17367 WOVd

[edAjodoy]
OTEE SHU “TBD ‘OD UlOW “geELs WOVA

“7 eyiwind snug

‘wey eyenoiued elanslja0y

"TN seplosssny esoutinses

‘spny wnjeqo] wnipidsy

“MG BotUOdef eIpPIeEMpOooM

‘J[Ney wWnrdtuwJopyeo umnipodAjog

Tyseyeye |
"I 2 oyes ssuoqeured

WOiIWNdadrdad
(*y wintwiselydiy) ‘zuery
(‘pAS) oelJaIne1[204

VuOSdOSSAN

‘JoIq eouodef
VITANGAVYOHLON

(‘A PBISOTIA) MOpAS
"H ® ‘d ({[ne4) eovisasoa

‘J ‘syeII] reseurqo}

(‘| Opelp) °F *syeITH
(‘191q) BjNdsntAde]

*P,juOd BUISaTTIA

18

[addy jo ued] LOI HH ‘88L7¢ WOVA
[adAjosy] ZL “MIOH ‘E8S801 WOVA

[JeA jo
ad AjofoH] “Te 39 EPPS JOPI2D “1TOELbl WOVG

[UOTDa]]09
ad41] WOV Ul pore Ssoulved “q190H

[adAjosy] 9€¢ “JOH “S789L WOVG

peploose1
you PI Inq [edAjosy] [Z7¢ “MIOH ‘7789L WOVG

[adAy jo ued] HH X2° 9777E WOVA
[addy jo ued] 682 O84D “‘HOIW x? 078 WOVG

JOM oeWeINSJeU eIIIUIIOg

“YosiIy epedasosajoy ep[NUa}0g

*|purT] Sluejnoioe esoy

‘gpAy ("DP Bsoonny eyUaog

“Musg sisuoyinb euljawwiod

"J BsO[NpuR]s vo[eJosg
IBYVN eIOYIsIAW eNaYy”

"sang SISUOUIS e[eIP9D

‘J “SyeIIPZ WinueO}!
‘Wun. asus;eWIa}eNs
QJIAes 9]8910q-1A0U

"IBA “JOOIYOS DULIOJISNY
Jeaysg lWoslopue

WNIdINS Vad

“MJOH 2 “SOC 190)

“MOH 2 “Syorr evoyesosd
‘J ‘syed oene7ya
‘wns eUROSYS

VuOSsSdOAVHd

19

[adAy jo wed] ‘u's ‘“MjoH ‘psgs0l WOVd

[adAjosq] 61692 2 978 WOVG

[adAy, Jo wed] HH xe ‘ouesny *s ‘osz7ze WOVd

[edAy, Jo ued] 1 16h 3ucT ‘p69, WOVa

[‘dds Jo adAjosq] pzz ‘sxq
"Ixeg “OOAW “PEZ1I9 WOVA

Cy wintydojryduy
St) “Dd wWnuosexsy untuao000yIIg

asung SIsusUIYS eIDeISIg

“OONZ 2 “Qolg eIpULLIAW eUISOI]ay

Aevig ‘Vy esojnpur]3 enasinosa

"YOY eUUTDUOD e]]NUDI0g

(‘eB elutsong)
“yy (MIOH
% “y2IG) Wydoprydure

WhidOdSOud

(aM 7 le], oetoejsid
‘d =) ‘wuing oevisuswayjs

VIAVIOUTId
(‘Ww elosdoyeyg) ‘wey
3 “WUWIND (‘sny) seUISOI;oOUr
WT THdOSAHd
(@ ey[901I[)
*pAs (Su07]) BeuTUINDe
SIXACOWOSVUHd
(‘zuel] oje010q ‘gq =)
UOSI9}9g ISUSSUTWIOAM

‘dss ‘pAg ovisaat

*p,jU0D WINIpIUIse1yg

20

[adAZ] €191 “MIOH ‘7p69L WOVA

[adA J) Zp61 “MIOH ‘1P69L WOVA

[adAj0s]] pOr “IOH ‘P6S801 WOVA
[adAq] ZZL1 “MIOH ‘Ob9SL WOVG
[ad41] €9¢1 “MICH ‘6€69L WOVA
[ed 1] 8961 ““IOH ‘088LL WOVd
[ad qr] IpSl “MIOH “L¢69L WOVG

[adAjosq] Soe “MIOH ‘8SSsOI = 6~SLL WOVA
[ad4 1] 8061 ““IOH “9€69L WOVG
[adAjos]] I[€EI “MIOH “SE69L WOVA

[adAjosy] poet “MiOH “€£69L WOVG

[edAjos]q] 1661 ‘MICH “0669L WOVG

slaty wingesojsod wumnypAydozng

avoorluoUusig

‘weyD 7% ‘TyOS eeydaooAw eiddry
"WINYOS eq[e eWOdd |,

“Od einpytu erpun’y

[Saig (‘TI9A) BISNUSA BISI}SOIA

‘ds wintus}d090uI

‘Jq Wnyesinu wntus}D099TII

‘ds eipuny]

“mINYdS 2 ‘Ing UINdads WNNseUeID

Cod
Toyouryg “Y Jvou) eoepiqeliy

ovaovauousig

“M]OH
ap syoer mypAydozns

“MOH
>? “syors WINjeININeI

(‘d erutsong) “wuind
(‘MJOH 2 ‘Wy) wnuseuLsd

“MJOH 2 ‘spore wnyewyed
"MOH “syoRe oerpuny
“MOH 2 “syoes wnytodunt
“syoer IABM[OY

(‘p erursong) “wun
(‘MIO 2 ‘Wy) sualjedap

"WIND sUIOJIWeAD
“MOH 2 “SyoeL WNseUsID
“MOH

wy ‘yors ovaepiqeie

“MJOP] 2 “syoes Winyewoue

*p,juod untpodso1g

21

[addy x9] ZELE “MIO ‘pZ6801 WOVA

[adAj0sq] [pz “MOH ‘I918L WOVG

[adAjos]Z] 6681 ABW ‘JJOpsyng
os[e ‘CpoLL WOVG [edAjos]] pegl ‘AeW-idy
‘JJopsyngs ‘WOV Ul 6€1 ‘PAQ ‘Wy “N “Weg

[edAj0s]] WOVAa
Ul 6 ‘ON I8unJ uRIXa] S,o/3U1Ig “87p06 WOVA

[adA4j0s]] WOVC UI LLZI “Pein ‘pds
[edAj0s]] O08 ‘“MIOH ‘TIZ8L WOVa
[adAjosq] c6se “MOH ‘998801 WOVa

[adAy, Jo wed] wizzesedg ‘79/8, WOVd

‘bores suaquinsep eueryduron

IvIIUYJUCICUIY

‘ds uinpsodsomJON

BWIWIS)SO[IYDIGG SB JUIIH
(-qstjeg) eyjayoind vaeiporg

aBaoeNV

gsoy WnaIne WNWsWeNs]
asoy susquinsold elonsne
SOON SNPIOSIA eITTONY

‘ds sninuesisiuy

aeaoeyUEIYy

"Iq eaovlyese * | = eade[eAO eUIODOT,

‘yy eiodsisaqo

"MOH 2 ‘SAL Iprodssoyjou

; "MOH
Ww “Jol, PeUIWA}sO;aYoIp

"WAdS IIOWeI}9}

*pAS eueresys

oy 3? syoer ereaoeid
“MOH 29 old Mpuesisiue
VINIDONd

(71 eruIsoNg) “MJOH 2%
“syour (‘dadg) ejooru0sa}

*p,juos umnipodsoig

pap

[jerzayeu adA ZT] €9¢ “MIOH “S6~8L WOVA

[‘u,Jjoo edAy] L9b801 WOVA

[dss
jo adAjojoxy] “Te 19 $9617 JOPIED “978901 WOVG

[jersayeur adA J] O19 “MIOH “TPI8L WOVG

[adAojoF] “WEA 2 9E9TZ aTtaes ‘pscrps WOVA
[addy Jo ued] ddd x9 1S6hS WOVA

[edAj0doJ] S79LL WOVA
[adAjos]] 79 “MIOH ‘S9SLL WOVA
[dss Jo adAjojoH]] OVA xX? 8SZ7LOI WOVG

["u,]]109
adA1] WOV U 9€Z1 “PAIN “WY “N “Wed

fadAy] 79S “MIOH “STL801 WOVG

Joneg ezJopURX eloBoelIY
“SION BIeIeIIp eoljasuy

‘Wy
2 ‘YOOH Sisus|iyd eziyOWsSO

‘ds eziyJIOWsO

‘dS
(J) esusulys umnuTjasolu0D

‘J ‘yooH louow eypAydioy

“WN (XY) esoging wuasig
"JT WNgje[Ndew WNlIOD

u
® °D (-YOoH) eurpurqosay elxAJord

DF

‘T (yoo) snuputqese) sniaydow AD

avaovldy

BLIeUOg

avaoepmAIewy

“MOH 2 ‘syoer eunusdal
“MJOH e1owOoIOd

QIIAes aeyopIed “dss
TyOu (-S) eefjourdund

“MJOH 2 “Syoer ejnpunu

QIAvS euRl[ausI3s
‘uund TweyeIs

(‘2 snpojly)
‘yy (UOC) seIUS3IIa

“MOP 2 “SYORL SIOISIP

aflaeg sisuauo3aso “dss

“MOP 29 “JIC WeidourAo

“MOH 27 “YY Joyjed

*‘p,juos elmldong

Zi

[jetsa}eu
adkT] 9681 APW pz ‘Jopeey ‘8IZ7ZL WOVa

[adAjos]] ¢€99 “MIOH ‘ELILL WOVG
[adAjereg] “MOH ‘PO0LL WOVG
[adAjosq] 6PIT “MIOH ‘O00LL WOVG

[jeriayeul odA 7)
WOVd U! 1917 ‘pain “PAS = ‘pLILL WOVd

[[elioyeur
adAT] ANd X2% UeD “qI9H x9 ‘1469, WOVA

[jeliayewr adAy (]
6 Sisualfiseig ‘ooAW ‘31 ‘€778L WOVG

[[etwoyeur ad] 9¢¢ “MIOH ‘1h08L WOVA

“SSOT SlIeIIo oWOsDAYeIG

“‘Wuog eazieds slieyoorg
"S19q (WT) SIAIOUIN) slIeyoorg
‘ds suueyooeg

98919)SV /9B998.19}SY

“JT B1IO}OUN stUaIpUY

“ASW, Sinesigye eay[iyoy

dvapIWMIyUY /avIde.19}SY

‘yNYIS Wsyueq winjejedAxQ

avaoepeidapasy

Aqsny twopuoul eajiAspueyy

avaseuAs0dy

sud; Yo sowooAyorsq

“MOH 2 “Syore
voyeds-sipieyoorq

“MOH 29 “SxOCL eye
“MJOH % ‘sour ejnqye

yeqng euellsajumnaeq

"OAD seaypIyoe

‘uusH IRIadAxo

“MOH

w “syoee sea]! Aopueul

*p,juod eluIdong

24

[edAjosq] €99 “MIOH *6L78L WOVA

[adAj0sq] L6L “MICH “SL78L WOV
[adAj0s]] [1 “MIOH “9978L WOVG
[adAjosq] €O1 ““IOH “889801 WOVG
[adAjos]] £16 “MIOH “968LL WOVG
[adAjosy] LEST “MIOH “€68LL WOVG

[adAjosy] p81 ““IOH “168LL WOVG
[edAjos]q] OLI ““IOH “9L8LL WOVG
[adAjos]] 9p¢S “MIOH “€L8LL WOVG

[adAjosy] 6p “MICH ‘O18LL WOVA

[rerzayewr ad] 1€8 JO 178 “MIOH “6P9LL WOV
[adAjoj0H]] 988h AAS “SEE90I NOVA

[terse ed J] OLTT “MIOH “9SPLL WOVG
[retajew edz] OF6 “MIOH “pSpLL WOVG

‘ds sireysorg

‘ds sireysorg

[YA BUSpI) slreysoeg
"Ssa] BIjopIouR] slueyooeg
“SIH Sisusjepulsy sireyoorg

‘ds stueysorg

‘ds sireysorg
‘Iq e[ewiour sieysoeg

"s1aq SUSpUBDS sIJeyooRG

‘PPaAA SISUSTAT[OG sNUIeTeI}OIN2H
‘ds sieyoorg

QuaaIy (ysing) snulseJed uo1es1Igq
‘qq syeAZso steyoorg

Aqsny ewuoiyes “Iu sizeyooeg

*P,ju0d 9B919}SY /VBIIEIIISY

“MOH 2 ‘syorr Bsnusasod

“MOH
aw ‘syorr sijiqeaidsiod

“MIOH 2 ‘syoes euooutsed
“syoer edo I9}UI

*MJOH 2 ‘syoer eurdout
“MOH 2p ‘Syoes Byesepul

“MOH
2 “syorr eysodusoout

“MIOH 2 “SyoRL es1sd0IduN
“MOH 2 “syoes enjodun

“MJOH
aw ‘syoer IWeyeEIPOIJII}9Y

“MJOH 2 ‘syors Banodxe
QIIABS SIUOISINIXa
“MJOH 2 “syoes BINsuod

“MJOH 2 “SYORL Bansuod

“p,juos eupsong

25

[on ul edAjos]
xa] ‘u's “mjoH “(ATUO aplys) 966811 WOVd

[adAjojox] “399,49 wnsesodwo0s
‘d S® WOV UI €¢7Z I3uny “Wy “N “Aq 2 “II

[adAjos]]}
WOVG "! p9S1 ‘pain ‘PAS = L969L WOVG

[adAjosq] 9975 “MOH ‘L10601 WOVG
[adAjosy] p79 “MIOH “18S8L WOVG

[-1ea Jo adAjosy] OT [-€9 UsuUaH ‘Eplpl] WOVA
[adAjosy] 1671 “MIOH ‘S9S8L WOVA

[ad Jo ys1ed] OETT J9A0[D ‘86E~7 WOVA
[adAjos]] Ce “MIOH ‘90P8L WOVAG

IMOIQ SNIIUD se
yeneg (‘psuoD) wnuoson, wnisd

“suoids (‘[YyNJA) JO[OIsIp uNISsIIZ

*ssecd (J) eureoe UOWOUDAg

daNpjBD/2vIde.19)SY

"Jd susdsapsos sieysoeg

“ds stieyoorg

aUuIdIH Xo[duNs uUOINsIIq

‘Iq BWOpAxo Jesu steyooeg

"slog BSOUTINIS slIByooRg

‘ds sireysoeg

*P,JUOD IBII}ISY /9BIIV19jSY

“MJOH
W Iq seouJOJIeS

QIIABS WNIOUTISSIe

*pAS ovuseoe

‘yy eouoyds
“MJOH 2 ‘syoee vyesqayes

usuusH eyoi0jd9]
“IeA “WUIND 2 ‘YY essni

"MJOH 2 “syous BlIvJopny

(podurejou
"qd =) suey ovjndi

“MOH 2 “swore eynooesd

*p,juod eBluIdoNg

26

["1eA Jo
adA,ooH]] Uapamg ‘WisysJeseT “D “677S7 WOVA

[‘sea Jo adAjofoHq]
puruly ‘uouleAoy “H 2 “T ‘116SL WOVd

[adAy, Jo ued] ZpZZ OPYD “68LEh WOVA

[adAjosy] avamnejues *q
se WOVA Ul gccl PAN ‘Wy 'N “UNE

[sea Jo adAjofoH] OELE “AIOH “ANd U!

[IBA
Jo adAjojox]] sueljes 2 sIoUU0D ‘18s WOVd

[zea Jo adAjojoH] JOpuNZ “p6ELL WOVA

[sea Jo adAjopoHy] OOS Aes “TSOEET NOVA
[zea Jo adAjofoH] J01S0.4 “890hp1 WOVA

[edAjos]] WOVG
ul {9L] ‘doing isunj “qey = Sp87t1 NOVA

[ea Jo adAjojoHy] ADjeAOY “L167El WOVA

‘doog (‘J) ansnyed winisitD

IH (CD winyyAydosayoy wintss1D

‘ds voinssnes

‘NN eURdTIOWe voINeUID

‘ds wintsid

‘yy (‘qpAy) HueWIpoy Wns

‘YNN WINIPOJIAIIG WNISIID

‘NN egNpa wnisD

"JT snueAd voInejUsD

edd “eA “T snuevAd eoineUs_D

‘we’] ESNYJIP vomneUID

*p,juod dvaNpieD/avI9e19}SV

QIAeg slysnjed “1eA

ayiaeg eouuay
“IVA ‘Qlase’] MYOse]

“wuIns euRIssueMy

“syour eyisinbo

opIAesS BURSIXOUl “IBA

QIIABS NUURUIpOYy “JeA
IIARS PIJOJIADIQ “JA

giaeg elladsayoo10q
‘ea “pAS esnjoul

QIIABS STAQTQNS “IBA

*sseg meds

a[IABS
esnyip “IRA “Od oeoINnejUIs

*p,juod eluIDoNg

27

[adAjosq] coze ‘syoer ‘qlaH “xo gcoezZ WOVA

[rea Jo adAjosqy]
“TRS “euapesed ‘Aysee[DoW ‘96801 WOVd

[ad4j}0aN] WOV Ul Opp I8ung ‘wy ‘N “SIT
[edAjos]] 1ase14 ‘S6LZ71 WOVd

[ad4jodo.] Wova
Ul CC] “Sxa “Joxa I3uny ‘“~pAS = E€7LL WOW

[adAjojoFy] “12
39 COOPT JOpleD “Woesary “gq se 996¢¢1 WOVG

[adAj0s]] WOVa
Ul OPIZ “AlUQ “OA ‘*WeNnyL, = pE98/ WOVG

["1eA Jo adAjofOH] “1eD ‘O16ZEI WOVA
[adAjojoH] WOVG W (e)1Z¢ “Suny I3uny ‘Yury

snjejsey snjeqen se
“Nad ‘q (‘yooH) eye sapueusig

ABI “VY eovlIoyold eliowouerydays

“‘yNN esourds eiusaposh7Z

“XY, Wnqeos WINIoRIOI

‘J “YOOH erjojrusjdse eoeuney]

QUuaaIH (*YOOH) evoenueIne silasosy

IVILIOYIIT/IvII8.19}SY

‘ds ejmeles

*sdor
(auseIH) wnyyAydospAy wns

"HI (J °T) wnououued wnisng

*P,jJUOD BBIINPIVD/IvIIV19jSY

“syoer eyesodsut

(ellowourydays *g)
sejoulieg (“pAg)
oevliauoueydays “IBA

IssouyIey “eA
8ZIA HSsouyIeYy

‘yy Lease

“pAg lapng

[EARS
W sojouleg siplsasose

‘Won, oejNelos

OARS eyIydoJOX *IeA

ajtaes roruouued *1eA

"p,juoo erupong

28

[-1ea Jo adAjos]] CIO “MIOH “8608L WOVG

[rea Jo adAjyereg] CgI-19 ‘wuiNnd “6LS901 WOVd
[adAjosq] WOVC U! OST ‘PAN ‘Wy °N “4queg

[sedAjosq] WOVd
Ul TPZ] “PAN “Wy "N “Yeg pue pe9LL WOVG

[‘u,]1[09 adA J] 86 “AIOH ‘TI9801 WOVA

[addy jo wed]

sour “AW “1109 27 ONO] ‘ajep oUeS

m0q ‘COI8E UNd Xe (ATUO aplis) 7P68TT
WOVG put SAN X@ “(AUO sa}ou) 67007 WOVG

[adAyuXs] SAN X92 (AJUO aplys) pS007 WOVG
[adAjosy] WOVA U! 097 “19D “A “WUE

[adAy, 21qeqQ01d]
COSPE UNd Xe (AJUo apis) €877LT WOVA

[edAjos}] WOVG U! CEPI “Alu “OoAT “WaNy]

“OC etoyoelaippnq eux

Aeig “y WUOUWWID] eI]]ayxOIIg

‘9q sediasiq wniojedng

Kei “y wniesoutdse wintiojedng
‘suIqoYy WnuenoTeW Wntoj}edng

avatIojedny/Ivade19}SY

uounmxoiy se ‘ds siissosy
uoUNTXOI], se “ds slJasosy

"ssa‘] (‘ssaJ) seasoi snddedoseurg

snjeqen se ‘ds somueuslg
‘WUag (‘sSa7]) BOLIIGIs BONO]

*P,JUOD IBIIOYI/9BIIC19jSY

“MOH 2 “syoRL ovueyIU

"Je 19 “WUIND eISNgol
“IeA “MYOS SeIUYNA

*MJOH 2 “321q sodiueut

“MOP 27 ‘301q Wnsesouldsa

“MJOH 2 “syour ejniodiseq

"y¥q sUOUNTXOy

“Aq 79 “Ia Wsopsyns
‘pkg tddedoseurd

“WUD 7 “Ad BINdIQIo

‘wony, SISUssnUIUI

*p,juod BIUIDONg

29

[adAjosy] WOVC U! 8IT ‘Pan ‘Wy ‘N “yueg

[adAjosy] 6€7 “MIOH “S659 WOVA

[adAjopoxy Woy] LZ7L9€
UAd x9 cEgE swesqy ‘(AjUO aplys) 116811 WOVA

[adAjereg] Eg0€ “sxoeL ‘879LL WOVA

[adAjos]] L¢¢ “MOH ‘9€Z801 WOVA
[adAy, x2] 989¢ “MIOH ‘886801 NOVG

[adAjosq] g08 “MOH ‘Sz7E8L WOVA

[adAy,
Jo wed] (pg ‘seig “q12H) 219 ‘S778L WOVd
BaSOI “gq SB

[edA4josy] WOV UI L¢ ‘pan ‘wy ‘N ‘‘Wueg
pue [adAje1eg] Z0ZS “MIOH ‘€68801 WOVG

“MAH Seprowpuerjsy esoinst A
OPA
(Ye]Og) SISUSDIIEISOD BIUDUIXITZ

aye[g (AeIn “Vy) eplosia vaelay

IVIYIULI]I}Y{/VVIIV.19}SY

“QpAy

COM wnyyAydoone] wnyyAydoug

IVIIUI]IFT/9VIIV.19}SY

‘ueg WnJoyI{nqm wniuojedng
ABI “VY eI[OJOVOIUOIOA eITJayOUIG

*qoy (‘JawWH) euetAntod eiionbig

‘yeg wmnijojisuojgo wniojedngq

‘borer wnapioyap wniiojedng

*p,jU0D dvallojednq/aVI0e.19}SY

“MOH 27 “301d eidnige
"MIOH 2 “syoes Bdoisqe

sojoweg liswesqe

*syoer myyAydorsa

“MIOH 2 ‘syoer sodpiyos
“MIOH 2 ‘Iq esIowsesd

“MIOH 2 “syour oeronbid
‘JaIq elodsAyoed

e
19 ‘WUIND vedasIsIago

‘p,quo erujsong

30

[adAjosq] 9L9 “MIOH “OTZTLL WOVG

[ad 4,007]
Zyry9E UNd Xe (Ajuo opis) 998771 WOVA

[adAyo3997q] “:09dS91 OIOLE FP 8PSLE
wWNd x2 (AUo sapl|s) pO6PZI SLEPT WOVG

[adAjos]]
WOVG U! 67ST “PAIN “Wy “N “4queg
= OILE “MIOH ‘(ATUO opts) TL8p71 WOVA

[edAy,
woy] ELSpE UNd Xe (A[UO opis) 716811 WOVA

[adAjos]]
WOVG U! Ist ‘Pen “PAS “L0681T WOVG

fadAy jo ued]
LI6GEE UNd Xe (AJUO apis) 906h71 WOVG

[addy x2] 7S “MIOH *7869L WOVA

[wy (‘3eds)
nuayed *g Jo adAJ] 6PE “MIOH ‘T¥78L WOVG

"WIg BURIAITOG eaEPIAO
"U19,J ([eWING) BIQedS “IBA

yams (J) Saplowpuerey sisdojeH

“NW eyAydosoeu ezryJoWesleg

‘INUdg WnsojUSWO) WNy]AYdiuIxy
"WUddIH
wW “suIqoY eIOJlOURIUOU BUISIqIOA,

"WUseIH
wW “SUIQOY BIVQOII PUISOqIOA
ayeIg (NN) BIOYH[NW eons A

"JSWIOH] (‘OQ) euedixoul sisdoa10D

“7 snioydosajysAy wntueyyeg

*P,JUOD BVIqIULTI}]{/ICI.19}SY

“MIOP 2 “SYOVL BURTATTOG

‘yy eurisajeq

"Yq seziyIowEs]eq

“yy mpAydiurxe

gojouLleg esolodiy) ‘IBA

“pAS stulje

(aeljaoue *‘q =)
“pAS “H ® ‘qd suejnwee

(aeyj09[9 “qd =)

"MOH 29 “SxeL eLONIppe
do [ULIed
Rjooluayped “IBA

‘pquos erurong

oT

[rea Jo adAjojoFy]

6968h UNd Xe (Ajuo aprjs) 606811 NOVA
[rea Jo adAjosy]}

€866r ANd x9 (AJUO oj0yd) O168I1T WOVG

[satseds Jo adAjosy]}
OS Bu0T ‘OCSrE UNd Xe (AJUO apl|s) O88>Z1 WOVA

[eddy jo
ued] TI€-19 ‘WUIND ‘T9EE9 ANd X° 978pyI WOVA

[sea Jo adAjosy] TOES

"MIOH “OS6EE UNd Xe (AJUO 9plfs) E1681 WOVA
[adAjos]]
WOVG U! 0€6 ‘PAIN ‘Wy ‘N ‘“Yyeg pue

[adAT] 9Z1¢ “MIOH ‘OP6EE ANd X?° EL8~Z1 WOVA

; [adAjo0s]] WOVd
Ul CIZI ‘Pan ‘Wy ‘N “Weg ‘g0681I WOVd

[adAjosq] 8619 ANd X° 1€6971 WOVG

[adAjos]]
G9g “Sxq “30xq “y “PAS = LL8bZ71 NOVA

*[SWIOH] Waselj euIsaqloA

"MH Sisusoeqin) eulsoqioA,

"J BPOIUISIIA BUISAQIOA

OyeI eqojousjs v1ain3iA

"WYOOTYIS Tyotyoayv9ez vajes

"OC CHAD eetfoyaeonin eae

*[SWIOH] XO *J “YOOH
2 ‘YUog SIMoJovloppnq esaINsIA,

oudeg (“1JO]) vaployap eisoiquiy

‘IjUeg Saploutseqioa snddedoig

°P,JUOD svayIULTIF{/IvI9v.19j}SY

So]dULIeg Laselj “IBA

do[ouLeg

BIENUIYOS “IEA
ByeUsOD “IBA “PAS BIeUSOD
soJoWeg sedeIO[YO

do[oUuLIeg
QBOBABUTOND “IBA

"yy ovayeo

sojouleg aelieonueyles

dIJOULILd SISUVDIOGED

*pAS ejayAyormq

*p,juod eIUDoNg

32

naoeue) “gq Se WOVC pue und u! [ed4q)
LSz Wun “wy sa “YU = 896971 WOVAG

[adAjosq] POLE ““IOH “176801 WOVG
“MOH
2? ‘JIC seIWO;oUWAS *g se [sedAjyereq] WOVG
Ul €Z6I ‘ZZ6T ‘IZ6T “PAN “PAS ‘[adAL]
LI6GEE UNd XA (Ajuo eptts) 906~71 WOVG
[sad A}0s]]}
WOVG " 0¢Z1 “PAIN “WY “N ““GHeg
pue O€9LE ANd X2 (Ajuo epl[s) 86871 WOVA
[ad Ajosy]]}
8h66r UNd x2 (AjuO eplys) OLOITT WOVG

[sadAjos]] SI p-19 “wuND ‘y0edsel
ZOCED pur E168S UNd = L8vII “y96ST1 WOVA

[sad Ajo0s]]
WOVG U 6€¢1 ‘PAN “Wy “N “queg pure 799¢
“MOH ‘pZ9TP UNd Xe (AjuO apis) 8L8P71 WOVA

[sod Ajos]]
WOVG U! OPS ‘PAIN “Wy "N “ye pue coce
“MIOH ‘18P9E UNd Xe (ATUO eplls) pr8y71 NOVA

"B BLIQSUBL] SP
gukeg (‘ABD) SOplolsolquie vISOIqUIV

"JC PIOJISIOAIP BUISIQIOA

‘UW RIWIOJOUWAD Se
ayejg (VNN) eJoyHiNw BIIINGIA

PURDIXOU BI[IDU
Se oye (ABD) BPNao} BISWITS

ABI “WY PI[OJIAIG PIUSWIXIZ
Avie) “Y RIOJIAIIG eIUDWIX9Z

Io9|e3
“3 se ‘Od eonnu sisdos10D

‘MUeg WNjeAO WNIpopueWssd

*p,jUuO0D VVIIULI]IH{/IVIIC.19}SYV

*‘pAg ovliosuel

“MJOH 2 “321d XOIIy

(suejnuloe *q) sojoulied
(‘pAg) suejnusee “JBA

QeI[IOUD “IBA
*MJOH 2 “31 sel[I0uU9
go[auLeg eISNQOI “IBA

sojouled

esoJodissaidap *JeA

9e1}09]9 “IA
“MOH 2 “IC VeNdeI2

“MOH
9 “oI Wpowpueulsep

*p,juos eIUIDINg

33

[ad&y, x2] €69 “MICH “€89801 WOVG

[sodAjos]] g6rre
UNd = 9SE “MIOH ‘OL9BOI Pur Z7P8LL WOVA

[ad 4}0399'7]
Hd Ul €767 “OU “10q ‘Wy UI ‘sun “‘UAS
WOl Elppe UNd Xe (A[UO apljs) OSspZ1 WOVd

[edAZ) ZLOL J9119H
‘SOPhE UNd Xe (ATUO apis) 9¢g~ZI WOVA

[edAjos]] cpr9¢
“MIOH ‘PE6EE UNd X29 (ATUO apts) O98~Z1 WOVA

[adAjoj0H7] 109
"“MIOH ‘TZ9LE ANd x9 (AfUO optys) P1681 WOVA

[adAjosy] ETS “MIOH ‘976801 NOVA

[adAjosq] ETS
“MIOH ‘7197p UNd Xe (AfUO apiys) pL8P71 WOVA

[eddy]
vOLZZ [87] SINY “VW “88LIE Sd’T X® 6L6HEI WOVA

[adAjosq] OIL-€9 “wUND ‘Og99TI WOVG

OAP[_ SHOVONI] CIUSWIXIZ

“YOOH
‘INueg (‘IWUNg) ByeUTWINDe vaRpIZAC

"J SNsoJoqn) snipuelay

ABI) “YW BdUIOJIeD epjoueroy

“yOOH
wz? ‘uag esonxayqns emojouwAD

axe (NAD susdsajniy eIUsUWIX9Z

*|duog 2p ‘quinyy euesixoul ejorpseny

*[SWaH] snooejeda snddedoio

“UOIOIH USII[[IS eIOINSIA

AeIg “Y WYeIQsays elUsWIXIZ

°P,JUOD svIYIULTIF{/I9vI9e.19j}SY

‘MOH 2 ‘syoee eupneul

*syoer epnAemjoy

“AMYOS 1pUeLIY

“YY seljowpueryoy

‘yy oenmojouwA3

gojouleg sasuajeulajen3

“MOH 29 “391 ae[OIpsens

"YY esaztingoys

(sue[nuee
“IVA BeI[INUN “q =)
“bpury euersaryyis

gogjoulleg IMYyselqsarys

‘p,quoo eyutzong

34

[adAjos]] ZZIS “MIOH “1L6801 WOVd

[eddy]
SI9TP UNd Xe (Ajo epljs) S0O6h71 WOVG
[edAy
x2] LESLE ANd Xe (Ajuo opis) 798P71 WOVA

[[etojeu

adX 1] S60ZP UNd X2 (ATUO apts) posp71 WOVA
[edAy

x9] LSppe UNd Xe (Ajuo apl|s) 198p71 WOVG

[jelsayeus ad AT] 1691
IN ‘STE8A ANd Xe (ATuo opljs) 6L8P71 WOVA

[sod Ajos]]}
6P8e 19D ‘dA = ‘SSp “Pan “Wy "N =E€pSt
“MIOH ‘9Z9ZP UNd Xe (AjUo eplys) Lp8rp71 WOVA

[adA4y] 2-909€
“MIOH ‘89SPE UNd X2 (AJUO opts) 188P71 WOVA

[edAy jo
ued] pOE9e UNd Xe (ATO aptjs) ¢96971 WOVG

P
BOON SB “[SWIOH] SUaIdISep vadseseT

*Zuoidg (‘ARD) BIEIUSP eIBINSIA

‘Iq (‘7 susdsainy eIys0g

‘Od
(yory) WnyeoeAIp wnipodwuejayy

‘Jq suelo snyueyey

-Jq_BIEpsoogns euUIseqsaA

‘nusg ey[Aydosajoy oueydajsoy
"WUsaIH

Ww ‘sUIQOY eI]OJoURJUOU! eUISSqIOA

ysing Sle]Ixe eA]

*P,JUOD IBIYIULI]IH{/IVIIC.19jISYV

‘yy oeo00u
(soueydajsol “qd =)

“‘pAS “H 2 ‘d ewoueu

“MOH 7 VIC BOUL

“MJOH 2 “21q Upodurejeut

‘Wy sipesseul

‘JaIq SlieNso1I

“MJOH 2 “301 souvydsysor

"SyoUr BIL[OAUI

"Yq BIXIULIOIUT

*p,juod BIUIDINg

39

[adAJ] gcs “MIOH ‘98E8L WOVA
[adAy] 7097p UNd Xe (AlUO aplfs) Z7O6~Z1 WOVA

[edAy)] 6Z01 Joquioyy
‘06SZ UNd xe (AJuo opis) [¢gp71 WOVA

[[elssjyeur adAy] Opg

"MIO ‘S8P9E ANd Xe (AuO opis) 106~71 WOVA
[edAjos]] a[3u1Ig

‘0097 UNd xe (ATUO aptjs) 9,8p71 WOVA

[adAjuAg] pOO7P UNd x2

(AjUO apt[s) LO6PZI PU LIS “MIOH ‘TIL801 WOVa
[adAy] ‘u's

“MOH ‘(A[UO apl[s) CLpPE UNd X° 668h71 WOVA

[adA]] Vos
weaq ‘HOIW Xe (AjUO apts) 006771 WOVd

[HA X92 uoutioeds
adA J] 68S7p UNd X9 (AJUO apis) pOI9TZT WOVG

[jetsoyewl ody] 977
“MIOH “9661 UNd x2 (ATU aplys) OL8PZ1 WOVG

"Oq eieiqes eluwAlog

Aen “Y WNeUNsIe UINIUDIIeg

ABI “YW SIJOW soIUSyIeg
sadiacig v1a}dojON se ‘[SwdH

Caig’y) snioyiains snddedoi9

“SUIqOY sniposiusajyye snddedoio9

oye] SIUSISUI vIIeD

ZNYIS
(PIL) B1]OJIIJOURVD eIUDWIXIZ

*[SwsH
Cig’ y) snioyiaino snddedeoz

BSOIO} BoIUTY se ‘ds eIpeyZ

"Gin. (MS) BINsIIY v1a}doO}ON

*Pp,jUOD svayJULT[IF{/IVI9.19}SV

“MIOH 2 ‘syoer oeruwAjod

"yy tuoyed

"syoer saotustpied

"ysipe vjoorddedojo

"PAS “H ¥ ‘d !ddojo

(ipoduiejow ‘gq =)
“MOH 2 “Syoee eVUIPIO

(2eIUSUIX9Z

‘d =) “M[OH 2 “201 voedo

suleyy b)e]GO

“Aq 29 “II epnu

"yy 9eJo)}do}0u

*p,juod eluIdoNg

36

[adAjosq] 92€ “MIOH “8EL8L NOVA

[adAjosy] 78 “MIOH “P7L8L WOVA

[adAy, jo wed]
SsoUuxIeH “UNd x2 (Ajuo aptls) 996971 WOVA

[adAyosq] Z9L-€9 “WUIND “7E697I WOVE

[eddy]
Hd X® pLOZp UNd x2 (Ajuo eptis) L98p71 WOVA

[edAy]
OPI “MOH ‘UNd x2 (ATUO apis) 76871 WOVA

[adAjosy] 968h Sd’T X9 EL6PET WOVAG
[edAjosq] WOV U! 7987 ‘PAIN “Wy "N “UNE

[addy] a[surig
‘Ss x9 UNd x9 (ATUO opiys) EP8P71 WOVA

[adA303997]] LyZ
“MIOH ‘ZZOLE UNd X29 (Ajuo opts) 9p8P71 WOVA

[adAj0s]] 31-19 “wund “ANd X° OL6STI WOVG

axel [AeMjoy eBl]OpaA,

dyAeI_ I9sOl BIUTIDEIIONS

‘ds eajoousw AY

oukeg (Atig “Y) ePl]OJIpIOS eBIsoOIqUIY

“J wimperoyiy wNIYydyIS

BSONXIYJQNS eIWOJOUUIADH se oxeI_
(‘WwW 2 ‘ED wnjepi09 wintydajsousw AH

‘H
® ‘g (ABD) SAaploljsoua BUISIQIOA

"NN PUWTXeUT BIyDEqpNy

"WUsdIH
a ‘suIqoY la[3utId eOINsIA

aye (INA) susosainy eUSWIX9Z

"|SWa] SUaIdIsep vaosese’]

*P,JUOD BBsyULTa]]{/IV9dC19}SV

“MOH 29 WoerL epinbeqns
“MJOH

I ‘SsyoRL eIUNIIEIIOIS

aZiA suopualds

gojauWeg seJOUOS

“myog Hydts

“MOH 2 “SyORL BOWES

*Zadg euvayiyos

“yueg oeryooqpnd

‘pAS “H ® ‘d eoprojound

“MIOH 2 “syoer eqoid

gojouLieg esstulajoeld

‘p,juoo Brulsond

37

[adAq} ¢9
BUuOT ‘pI9bE UNd X92 (AjUO apts) 6S8pZI NOVA

[edAy, jo wed]
Hd X? 16ZIt UNd Xe (AJuo apts) 496971 WOVA

[adAjosq] 7781 “MICH ‘9068L WOVG

[adAyudg {] 1oAeWZ ‘S068L WOVA

[ad4j0anN]] WOVA U! 6967 ‘10D “4 “Weg
= (Aju apl|s) 8OSP€ ANd X29 pue ssgpZ7I WOVA

[addy xa]
“MIOH ‘O19%p UNd Xe (ATUO apifs) 698~>71 WOVA

[addy Jo weg] gp SuIpooH
‘OSSZp UNd xe (Ayuo aplfs) g98p71 WOVA

[adAjosy] “mjoH ‘99¢7p UNd x2
(A[Uo apis pue sydesZors1woj0yd) go6pZ71 NOVA

[adAjosq] ggpe “MIOH ‘7pL8L WOVG

“H
2 °q (‘ABD) Saploljaous euIsagi3 A,

‘ds winijuex

“OC eydaysoysin elapayy

"Od erydajsoysin erjapa yy

"SI@AA ("']) SI[RIUSPINDO eUISIGIDA

‘ds eulsagioA,

Aeln “Yy evoplojyop eiIn3IA
“pout? e1ogiqny °L

SB “SSB SIUIOJOeQR} EIUOTpI

aye[g (AeID “y) Wowyed sidajoskyy

*P,JUOD svaUBTIFT/IVIIv19}SY

BUO’] SPISOUOUIIX

"MYOS HIpUeX

“MOH
Ww “syour ejoonjapam

(ipoduejaut -g =)
JoAeW “q oeljapam

“MYO IBUISAGIOA

“syoer BPA

"syoer Sodipising

(aeljaoue
“IVA OeI[Z0US *g =)
“MOH 29 “21 oeruoIPH

“MIOH 29 “J0IQ] BSOgo|sqns

"p.quoo erutong

38

[adAjosq] Zp “MIOH “87801 WOVG
[adAjosq] LOLI ““IOH “97P8L WOVA
[adAj0s]] Cost ‘MICH ‘S7P8L WOVG

[adAjosq] £79 “M1OH “9€08L WOVG

[uonsajjoo adAy) Ao. 27 ploy ‘96SLL WOVG

‘yeng ensnuy ‘TL “MjoH uou [adA30}99T]
"yEND POJOS ‘ZOT “MMIOH “SPL801 WOVG

[edAjos]] 1p6 “MIOH “ppL8L WOVG
[adAj0s]] 66 “MICH “EpSLL WOVA

[adA}0399']] § x9 (A[UO 9pl{s) ZL8P71 WOVA

[adAy, Jo ed] Zpes “MIOH ‘LE0601 WOVA

“wUsdIH eyAydAejd eydsesostyo¢
‘dig ‘yds stAlJourpronjjed o1sauag
‘dig ‘yos stasourpronyjad o1saues

‘ds o1d0U9S
"yOOH esojnpurys eIstwjaD

IVIUOIIIUIC/IBIIV1IISYV

‘Ig ‘q SUddSaINIJ SIXT],
axel elposmises esnny
"ssa‘] esosni eisune

JvISIIN|A/2VIIV1IjSY

Aeig “yw ejeydaoopod eiuswixeZ

‘ZY9S eulodse eluazn[eZ

*‘p,JUOd svayJULT]aH{/IVVde19}SY

“MOH
w ‘syorr aeydiesojstyos

“MOP 2 ‘Syoes esoranjoid
“MIO 2 ‘swore ejnssd0I1d
“MIOH 2 ‘syore epnosnfeu

‘uuNns sIsua}uOWIs9

‘YY syrxiy
“MIOP 2 “Syoee eqns
“MJOH 2 ‘swore BDaJop

“MIOH] 29 ‘JIC, VeIUSUIX9Z

"yy oeluoznyez

*p,juod elu1d9ng

39

[adAjosq} C971 “MIOH ‘OL8LL WOVA
[adAjosq] 67 “MIOH “998LL WOVG
[adAjereg] 1p “MIOH “819801 WOVG
[edAjereg] 1971 “MIOH ‘OOLLL WOVG
[adAjosq] 9171 “MIOH ‘S89LL WOVG
[adAjosq] 097 “MIOH “pp980I ¥ 89SLL WOVG
[adAjo0sq] 7961 “MIOH ‘OSSLL WOVA

[adAjos]] 6611 “MIOH ‘SOOLL WOVa

[adAjos]] PBI “MIOH ‘P669L WOVA

[edAjosy] WOVA U! 76:I
"pry ‘idAID avjuelg ‘WsqIT xe CEP9I I WOVd

[edAjeaeg] pS7¢ “MIOH ‘L00601 WOVG

‘ds vluoulaA

"S19q SOploidsios eIuOUIIA
"YAH PSO[NdsoyiN eIlUOUIIA
“ds eluoulsA

ssa] ey]Aydosoew eIUOUIOA
‘ssx] euviddap eiuoulsA

“ds eruoulaA

"dig ‘yos eryoInoiA3Ie eIlUOUIOA

"SSa7] ESNYIP eIUOUIZA

IVIIUOU.II A /9VIIC19jSY

"]aWD UsyoNy o1dauES

"Iq snipoyijnsue o1sauas

*P,JUOD IBIUOININIC/IBIIC1IIISY

“MOH
w? ‘syoer stjiqenodut

“MJOH 2 “SPOCL SIIQeyeyy!
"MJOH 2 “Syoes vauopl
"MOH 2 ‘syore Hepuny
“MJOH W ‘syoRL eysney
"MOH 2 SYOeL eIDIOSIP
"MJOH 29 ‘syous eauessdoap

“MOH
F “syoes sifiqepueyye

*M[OH
w? ‘syoer sljeuonuse

‘qQI'] stuolsouas

"IY elOo0I1N9UeS

*p,juOd BIUIDONg

40

[adAjodoy] 72-69 sutuunD “169671 WOVA
[adAjosq] OLSE “MIOH ‘TO7LL WOVG

[edAj0sq] 6811 “MICH “7S88L WOVd

[adAjos]] OZPI ““IOH “8788L WOVG
[adAjos]] 06h ““IOH “97L801 WOVA
[adAjosq] OS “MICH “61L801 WOVG

[edAjosq]
Z691 ‘Skid “GI9H xX? IN “61€8L WOVG

[adAj0sq] €€Z1 “MIOH “0878L NOVA
[adAj0sq] 1061 “MIOH “8L78L WOVG
[ad4jos]] gr1 “MICH ‘90L801 NOVA
[adAjosy] azuny “gp6LL WOVG
[adAjereg] 168 “MIOH ‘TS8p1? NOVA
[adAjosq] p9OI “MICH “€88LL WOVG

appeay (“Aue}) syloes3s eluoyeyy
"NYO eIOJII) SIIOQIOG

guaoepllieqieg

‘ds eluouloA

‘dig “y9s
(‘Oq) ededosp3io sisdowsojjiue,

"J ediesola] BIUOUIDA

“SMqGH eBsoTNIsoyIN BIUOUII A

"ssa’] sisuaje[d eluOUIOA

“ssa7] eljoyiisoddo eruoulaA
‘dig ‘yog Soplorsuassa] BlUOUISA
"Jq edivsoul] eIUOUISA

“UOIAIF WZUNY eIUOUIOA

“WAH susyed eruouloA

‘Jq_esolienbsqns eluoule A

*p,juoo IVIUOUIIA/ svIIeIIISY

‘wuND seTuOYyeUl

“JIC SeIOJIN-S119q10q

‘MIOH 2 “SYoey StIqetuaA

“MJOH
ap ‘syoer sipisdoulsoyuea

“MJOH 7 “Syoes Bed

“MOH 2 “syoeL eyeowsd

‘yaIq sinsuid

“MOH 2 “sxoeL sttrqused
“MIOH 2 SyeL eqsjod
“MIOH 2 “Syoes eyyjou
“MOH “sour 1IZ1uny

“MIOH 2 ‘“syoRE eyenbovul

“MOH 2 “SOL PISIAOIGUII

*p,juos vrulsong

41

[dss
Jo adAjopoHy] “12 39 PIIST JOpled ‘Se69p WOVA

[adAjojoH] BITZI APOD ‘7S8L11 WOVG

fadAy]

LST !sunJ ‘ued se Joye ‘pL6S17 WOVA

[adAjyosy] 68¢ “MICH “L6L8L WOVG

[adAye1eg] 979 “MIOH “199801 WOVA

[edAjosq] WOVC UI OPI “PAIN
‘WY °N “queg = ‘U's “MOH ‘9V9LL WOVG

[adAjos]] COE “MIOH ‘6PSLL WOVG

""T BILOJIPI[9q oulWepsIeD

10a
AINIC (QUddID) sI[vs10g BLYSMOTIWIS
‘ds siqeiy

IBIIVIISSVIG

"SAH esoulsiyny ensojoumnoy

ws
‘q wnuidAjesosKyd uinidonosyH

aVaVUIse1Og

*|duog 2p ‘quinyy sijjow ewiosa

‘q Wnyesinw wniuajs090TpIg

avadvIUOUsIg

QJIARS SI]eaJ0qG
‘dss ‘pny winsesajionio

aplaes 1Apoo

"AQ 7 MA Ppooiqere

“MJOH
w? “syoes seIOJAUINO}

"yuy
Ul “MOH 2 “WY eayis

CmunD (Aq 29 “1D
SUPULIOJSURT}

winipodsolg =)
*MIOH 2 ‘pAS esonixs

CP

wnipodsolg =) ‘yy ul
“MOH 29 “YY suojjedap

"p.quod erujsong

42

[adAjos]] p6L “MICH ‘PLSLL WOVA

[adAjos]]
WOVG Ul ILST ‘18S Sezolses “OID, “ISOA,

[adAjosy] eseurysox ‘7OSLL WOVG
[adAjosy] 7SZ “MIOH ‘79801 WOVd

[adAjosy] pe9 “AMIOH ‘SOPLL WOVG

[adAjos]] S19 Sed ‘8LL8L WOVA
[adAyooRy] OPLARS 2 LZLIT JOPIRD “P86r>y1 WOVA

[dss Jo adAj30j0H]] WOVC Ul
ZS9 “Sx ‘Ixeg ‘Ooh ‘UOYJOS = LP7I9 WOVA

[dss Jo adAjojoH] 61 IT PA1WUed “OSP16 WOVG

“VIR, BSO[MSY BIoWOd]

JBIIE|NA[OAUOD

“7 suadas eptydosdéy
‘qunyy, snowodef snipueiq

“PITTA, BIBAO PLIRIIAIS

avaoeAydoAied

"J vaURsI3s sWIO0I|D

avaoeplieddes

qusaIn “gq (Cully
2 ‘yooH) wnyyAydorse] winipodAjey

uoskeg bJa0uI equid

Aelgy “YW eIojipod sutwepse|D

"J RIOJIpI[Eq ouTUepse

*P,JUOD BBIIVIISSeIG

“MIOH 2 “SyoRL Bpuansunsip

yoyonly
Joke styuadas-epiydosdé3

“uudp] ‘qd eotuodel-1mpueip

“MJOH 27 “UY esuojop

"MOP 2 “SYOVL StU0a9

‘wuing WpodAjayp

QIIARS ejOoIQeIp

ajiaeg sisuasurwo0AM “dss

gojouled
a oplaeg eonoseau ‘dss

*‘p,juod elUDong

43

[adAjo}oH] WOV UI LOS 18unJ “]eD

[adAjos]é] $76 staeq ‘SLL06 WOVG

[adAjosy] eseutrysox ‘60€LL WOVa
[adAj3osq] 10AeWZW ‘p80LL WOVd

[adAj0s]] pOL0Z 2804 ‘ISOLL WOVG

[adAjos]]}

42d ‘WOV Ul srg “Alun, ‘ood ‘“uronY

[adAjereg] Cpes “MIOH ‘€L6801 WOVG

[edAje1eg] OLES “MIOH ‘696801 WOVa
[adAjosq] ‘u's “mjoH ‘1S6801 WOVG

“NLIg Nuopsuo0s sndio¢

noog evouInga xale9

sAYoeIsoOIO[Yo
"IBA ‘qunyy eotuodel xareap

[yea snsnyyip sniadA5
aevaoviadA7
‘ds elinsuy

avaIEVIGININD

“J sisuapued snujod

dvIIVUIOZ

asoy esojidoyuI eaowod]

'S 3 “Y seploonwunu esowod]

ssOY eULINOSTeM BIOWOd]

*P,jUOd dead" |NAJOAUOD

Q[IAvS TUOpsu0d
SIARG

"¢'¢ oeIpsoydays-saoes

‘JoIq seotuOdel-sio1e9

JoAey stsuatnbonue

"WUIND 2 “YY seLINsue

‘yung eMuesorAydiod

“MOH JO;OOND0U

(‘WE snpojTy) “IsuYyor 2
‘iy (U0LIQ) eIodsojesaur

“MOH SIUSISUI

*p,juod BIUIDINg

44

[adAjosq] snouihag ‘(69p8L=) 6£0L WOVA
[adAzoJoH]] “12 39 LOSTT JoPIeD “9176ET WOV
[adAjosy] pIpl “MICH “9188L WOVG

[adAj0sq] ¢gg “IOH “8918L WOVA

[adAjosy] POS “MICH “8S18L WOVG

[dss Jo adAy] €6967 altAeS “C06S01 WOVA

[adAyosy] eseurysox “SE6LL WOVA
[adAjosy] eseurysox “8p8LL WOVA

[adAjereg] OLLOE Add xX? C016 WOVA

[adAjos]] 806 ““IOH “€69LL WOVG
[adAjereg] Eppe “MICH ‘766801 WOVG

['Jea
jo adAJ] “Te 19 E8HET JOPIED “gz7e6el WOVA

[adAjosy] WOVC U! 88 “PAIN “Wy “N “WHE

"wey eorueayAsuad Xoled
"slog snuvoliowe sndio¢
‘yory “OT xesay snsadAD

‘ds verodsoyouAyy

wuny snueiaXeul snsadkD

wey] eotuejaseul XdIBD

JoyIeID
(‘noy) snunosas snjjsounf

‘ds xoied

‘ds viuyey

“yory “OT xesey sredAD

‘uJay eUeABMIOY SITAISLIQUITT

"J uoryorssAjod umnsoydig

“OJ, UZMUIAMYOS Xo1eD

“p,juos avaoeiedAD

‘wuns eorueyind

g[laeg stuesooA0so

“yy eINjoAqo

“MOH 2 “SyoRs e[NsuUo[qo
“MIOH 2 “SyoeL BAIOOU

IAS eNUsINe|
‘dss *zuviy, Bojorey

‘Jai, Wypeount

‘yaiq eurpeAYy
AaiZuiq eeruyes

“MJOH
a ‘SyoeL SUSITA-OARTJ

"YUY sIpiyAsiquily

glares lpissede
“rea ‘wanyy W0ydore

‘yy wedAs

*p.juos BlUsoNg

[adAjosy] WOV UI Cpl ‘Porn “Wy °N ‘“queg

[adAjos]] O¢p9 UOpsor 2 SyIeg ‘prELS WOVA

[adAjosy] J9119H ‘8p6Sh1 WOVG

[edA}0s]] 066 ““IOH ‘L89LL WOVd

[adAjosq] 0O9T “MICH ‘0869L WOVa

[adAjosq] eseurysox ‘(L6SLL=) 9Z7EEE WOVG

[adAjosq] 078 “MIOH ‘~19801 WOVA

"QaSIID (‘WIS) Bxa]Jop BIUsTey

IBIICUBI}UIS)

"112H (‘yuag) stjjour ejasoreg

avaoRqey

ushay BIeps0 eIquoydng

‘ds eiqroydngy

‘ds uowsjsounsy

avaoviqioydny

‘qunyy, suosund snusearyq

avovoeueseerly

*[SWOH] BIeLIosIONed efnesnes

svadEIUITIG

‘MOH 7 “WY eluayey

‘wUUND sejasoied

(‘A Opern)
"YUV (Ad 2 “IED Bela

“MJOH 2 “SOREL eyEISIJ

“MJOH
3? “syoue sluouIa}souTIIe

"JaIq XO ‘UIYSOX lUseRerIS

‘yy sipuvssioa

‘p,quod BruTDoNg

46

[adAj0s]] HH X9 ojouneyes “E687e WOVA
[adAjosq] €661 99UD “96LEP WOVG

[adAjosy] edeurysox “T1S8L WOVG

[adAjosy] uondiiosap

JeUIsIIO UI Rep Yew spoqe] [TV “"WOVd

Ul [1@ ZS ‘PAN “Wy “N “Ye™ ‘7pZe Hsun{
‘Wy (NAG IE S€SL 190 “A AD 2P WA

[adAjosy] 61 “MIOH ‘69SLL WOVG
[adAjosq] JOSaN ‘L7pLL WOVG

[adAjosy] ZLET “MICH ‘866LL WOVA

‘QONZ 2 “qalg Winsowlsses UNITAS]
‘ds stsdojAlog

aVIepoUIvUle HY

"wmIxe, Wnouodel saqry

"MOg (‘S19q) BSNde] Seqry

IVIICIIV[NSSOINH

‘[1Y¥q Hsusyoo wntuesey
EIJOD winuelssjJeq WNTURIED

avaouluelesy)

‘TYOS 2 “wWeYD smeE[Nounped
“IBA “URI, SURZIIO SNYULISIT

*p,JUOD svIdIVUBIVUIS)

“ysox
w? ‘J ‘syeIY 1ojoweyes

“wun sipisdojA109

‘uusH ‘dq eotuodel-isoqu

“MOH 2 “JIC aesoyIEd

“MIOH 2 “SAoeL eIUIISIP

JoBan sisuonbiyjes

“MOH 2 ‘SAOVL WVUrISH]

*p,juod erulsong

47

[adAjudg] ceg “MIOH ‘pgcsL WOVd

[sadAjereg] 67 “MOH
“88€8L WOV pure LZ¢ “MIOH “68€8L WOVd

[adAjo0sq] 699 “MIOH ‘OL78L WOVA
[adAjos]] LZ7S1 “MIOH ‘LS78L WOVG

[adAjos]] £67 “MOH ‘089801 WOVG

[adAjosq] ¢gz “MIOH “999801 WOVG

[adAjos]]
SOL “Bed “PY = 8g ‘069LL WOVA

[adAjereg] CEOS “MIOH “906801 NOVA
[edAjosq] 7/8 “MIOH “19PLL WOVG
[adAjos]] CZET “MIOH “S9ELL WOVA
[adAjudg] 188 “MIOH “pZ0LL WOVG
[adAjos]] 688 ‘“MIOH “6669L WOVA

[adAjosq] gce “MIOH ‘0669L WOVG

“SH eyeydasooney erayes

uo}INg Msasplid Ҥ pure
*blig stuepnotavopnasd erayes

‘Ijusg BeIOpO sndAH

‘usg wEpNoosey sudAY

[RD 2 “Ue saproydAy elayes

"luag eyueona] eyruND

"WUD HEINI[AIs sHdAH
‘Nog weunsed sndAyH

“ds erayes

“‘Itusg eroyiuidies sndAy
‘Ituog eyeydeoors sndAY

"yuag eselap eIAleS

‘bug eueazjuny erAles

dBIOBIUICT

“MIOH 2 “syore vues

"MOH
>? ‘syors vonasAydiod

“MJOH A ‘syoer eywossod
“MIOH 2 “syoer vyounziod
“Wy

Ul “MOH 2 suleyy eupodut

‘MIOH 79 “YU BIeOsTy

“Wy sinpy

“MJOH PO}SIP

‘MOH 2 “Sores eyeGINjUOD
"MOH A ‘SyoeL woweayes
"MJOH 3 “syorr eoyitdwue
"MOH 2 “SHEL eJooIq]e

“MOH
wW “syoer voIeUIZIUSe

*p,juod eluIsoNg

48

[adAjosy] extepnyo ‘Oo7TSEE WOVA

[adAjosq] ounsyey, “C6gze WOVG
[adAyosq] 2187 O84D ‘E9LEb WOVG
[adAyosy] 3967 92ND ‘POLEY NOVA
[adAyosq] 9261 92UD ‘E8LEry WOVA
[adAjosq] pZZ1 O24D ‘98LEp WOVA

[adAjosq] 398 09Y4D *LELE”V WOVA
[adAjosq] 1667 994D ‘66LEp WOVG
[adAyosq] O97 92D ‘TO8Eh NOVA

[adAjosq] C98 “MIOH “€1L8L WOVG

‘yegq eouodel eyplog

avooell]

IVJEN eIOJIZUO] sUydepoudy
‘ds wiozueg

‘ds vaslog

‘ds snqiyory

THA “a B1OsTUYDANS BOpurT
‘ds elopur]

‘ds wnwouwleuulsd

‘ds wlozueg

avaoBiNe’y]

‘yjuog ewEjnoIUed sjeoeyds

*p,juod svaoEIUEy]

"YSOX
"J “syed Testepnyo

(‘1 ajaysousX) “J “Sie
(‘J “syedi{) SIsuSUINyeAe}

"wuInZ eysodas
“WUIND SIULIOJLIIUWTOS
"WUIND IYSeUl

"Wun. BjOoTINe]
"wuInd eIodsouos09
"WUIND P[OOTWOWeUUID

“wUIND BI95

*MJOH
ap -syoer Bjoorjaovyds

“p,juos eyujond

S — [adAyojoyy] s0pqeo 2 18g ueuoyyNyY ‘7L9hL WOVG “quaysioy (“T) eunosas eIpAOTT OARS SISUSUOAANA

[edAjos]] ‘u's “MOH ‘808161 WOVad ysing suesaja snuope3sh7 “MIOH 2 “pAS esoumni3

[adAjosq] soyoniD ‘(TTT€8 =) ZOTLL WOVG “quayoioy (“T) eunoses eIpAO[T yoyonID “C lesseq
IVaVYULLIA]

[edAjosq] Aempoy ‘27€8L WOVA "YOOH Soplopis snwpuriseld “dyo~ Mpuerseyd
IBIIVAIC]

[adAjosq] TOvT ““IOH ‘OTE8L WOVa ‘ds siisidosajaH “MOH 2 “syoer eyernjoid

[adAjos]y] pEZI “MIOH ‘L68LL WOW “ssng slsodues eliojsiueg “MOH 2 “syorg b90IUI
aVaovlysidjeyy]

[adAj0sq] OSLE “MIOH ‘006801 NOVA ‘ds vaydnD "MJOH avoydno
avoovlgyAy

[adAjosy] uosulqoy ‘Z7108L WOVd “QalS Saplonsejad sny~uelOy] “dT yoW_ epoorpueisoy

avadeyjULIOT *p,juod eruidoNng

50

[adAjosy] “J “MeN “Oppee WOVA
[adAjosq] WOVC Ul 9€ET “AU ‘Oos “'WaNyL
[adAyuds] I7SLL WOVG

[adAj0s]] WOVA U! POE HUN “UY ISEM\ “WUD

[adAyosy] gge-79 “wuND “g6h901 WOVG
[edAj0sq] WOVA U! L8¢ I8uNy “WY Ise “HUD
[adAjos]q]

eieyy ‘(A[UO api[s) pE9ST UNd xX? O€8S8l WOVA

[sea Jo adAjosy] OEZ-19 “WUIND “E611 WOVG

[edAjosy] “WIV “667801 WOVA
[adAjereg] 0S9 ““I9H “66801 NOVA

‘Ig’y suodai snimday
"J snowizA s1uosodoipuy

‘Ides (‘Oq) smeId uosodoquihD

“LIO], Saplome snjoqosods

“OUOWH
C¥qH) wmequiy wNHaeYysosdig

-idny 2 ‘ull, BIeWOD edns

Bley yeyory xisAH

“"] stuotusospe epnsiiy

‘yorH
(-YOoH) eIop1Buo] eypLAoweElED

"gd 29 °H Seploryouss stuosodos0y

avaov0g

‘J syemy wmdsy
‘wony], eURISI[a
“ssey, stuosodoquiAd

(suelinxn]
RISO} *“q) DUIBIDH 7 “WUIND

(yyy) suelinxn] “JeA
“AT 29 “1g UpuerdAro

“WUIND sISUasOsIyS

su Weng

vlepy ovouodel-aejjoidse

ulesny 2 “WIND seIeJORYS
“yea AdelL, oepysie

‘yaIq euosiyduie

“MOH 7 ‘Wy stuosodosae

*p,juod eluidong

5a

[edAT] 88617 UNd X9 (AJUO sajou) E6~S7 NOVA

[rea Jo adAjofoFY] 6ZL MAS ‘Oppez WOVA

[rea Jo adAjojoHy] 16ZE aTAeS ‘OL0E WOVA

[adAjosy] edurysox ‘E9¢g2 WOVA

[dss Jo adAjof0F]] BQ9p BIARS ‘1TOZI6 WOVG
[adAjodoy] sure; ‘OLE~7 WOVA
[adAjosq] 926] °OU INIA ‘Lp6Sp1 WOVd

[adAjosq] 68¢ “sxq
"xeg “OA ““UIOS = ESbZ “UIOS ‘E1019 WOVG

[adAjosq] cge 0245 ‘oLLEp WOVA

[adAjosq] pRpT C24D ‘ZLLEp NOVA

"YOM PuRLaqinYy sNsOIsy

‘Iq CTyeM) eoindindone eapoje,

“Anvog (‘J) ByeIOpo I0],ydIJAIY

*YOOH stasoutipenb eiuyjog

“pueip WzyeYy bod
AaseA, (19310g) WUpIUIs BOITaWy

"Uli, SuatInid wnstueg
‘uqliog 1ajsutd edng

‘ds snsoiserq

‘ds eruljow

*p,jUOd avadv0g

stlovisovid *1eA

(-9 erutsong)
a[Aeg (2]1AeS)
HISJOUUOD “IBA

oplaeg eueljoqgero
‘JA ‘YY siloveisoeid

‘WIC
stAJoulpenb-serutyjod

gpaeg vonosedAy
‘dss ‘yO stjesowou-seod

Joyory evorxopesed
"AW 2 ‘[Iq sisuanyeo

‘unUND
3 9U9dIDH e}eU0IODOIU

‘wUINd eJ931I0UI
(IL61 “wund spy

“yal ejooruyserdip
‘d =) ‘wuing Pjoonutjour

"p,juoo elupsong

Oe,

[adAjo0s]] OZOE “MICH ‘¥Z0601 WOVd

[sadA4jos]] WOV U! 9811 ‘19D ‘A
““yueg pue ‘77¢¢ sung “Wy *N “Aq 2 “1d

[-1ea Jo adAjosy] WOV UI CLOS “10D !suny ‘ee

[adAjosq] WOV U! Br
JOD ‘dq pue Tple isuny “Wy *N “Aq % “Id

{ad4j0s]] WOVG UW
SLrE WsuNJ “WY *N “Aq 9 ‘II = 99SL6 WOVd

[adAjosq] 69-79 “wUIND “160911 WOVd
[edAjosq] €Zp-79 “WIND “799901 WOVG
[edAjosq¢] yoYONID *L098L WOVA

[adAjo0sqZ] 9681 “Idog OZ ‘“MIOH “710601 NOVA

[adAjos]]
WOVG U! 9p7Z !sunj “Wy “N “Aq 9 IID

[adMjodoyz ‘adAj0s]Z]
epemes 279 epewlex “HH Xe p6sce WOVA

‘[Soig BaWielsuo] epnsiy
*XYOIAY Winaoejes winjedseg

‘yory soprouswAy sisdozAIQ

‘ds uosodosA1y
se ul], e[npltA edng

soploiuesieds
X91eD SB “YT BeuNoed eunseds

‘Uap esojnuey epysiy
"[IYq SNIpOJIAsIq UOsOdOII]IS

"JT BIQNI voNIse.J

"SayINYIS 2 “Ws0Y SIQIIQUIT eIIR}IIS

snuexa) “S SB ‘Jal
(‘NN) smMe[Noiued snpreuUOpsyos

od
(-‘J) eoovUIpunIE sysoisewEled

°p,jU0D 9BI9"0g

sWUWOI]
9 “Wy ul “MjoH eorun

"queg 7 ‘lIq BeLsqns

‘wUIND 2
QUddI) SIpIsdozZAIO *IeA

SI]JJaysqns “IBA
“AS 7 ‘IA Stlzaisqns

“ye
® ‘Ilq soptoruesieds

BOIIOUOS “IPA
ulesny] 2 “WUIND ed1JOUOS

"wUuINs stuosodol9]9s
JOYONID seIQnJ-seI]19s
“MOH 29 ‘21 VelIEIES

“9[SUIMS
9 “119 Ipreuuopsys

OI] PULIOJISUeI

*p,jUOd eIUIDINg

53

[IBA
Jo adAjojoHy] atavg 2 sajaueg ‘11679 WOVd

[edAy, xo] sueagq 2 Aoely
‘(Ajuo oyoyd 2p apis) O1Z79E UNd x9 WOVA

[rea Jo adAjos]]
prT “WOW “xeg “oA ““YIOS “LOE1I9 WOVA

[adAy Jo wed] souor
“AW ‘(Ajuo o0yd 2 apis) ZPZ9E UNd X9 WOVA

[adAjos]]}
WOVG U! gggz Isunj “Wy “N ‘Aq 2 ‘Id

["1ea Jo adAjosq] QT Z-Z9 suMUND ‘pI I9TI WOVG

[adAjos]] WOV U! 8p ‘10D “A “Ad 7 “IA

[adAjosy] WOVC U! [COT !8uny ‘Wy ‘N “IIE

“] eyeOLIeAIP XO[Yd

suosund eleusry
SB ‘NN eIOJIsUO] XOTYd

Ryeigrys
"d se “‘Uag Hpooy xojYd
"YOoH Hsejsnop xojyd
IBIIVIUOUII[Og

ysen (J) sueinu
uinnseysiog = “J wMesA wnotueg

Aase A tAazyonq snjogosods

puny (“xyotpy) Jodse snjoqosods

"1IOL
(*xYoljA]) e[npusdiino enojainog

*p,juOd avadv0g

aajouledg
BILOLIVAIP “IBA

SIISBIJ “IBA
[eH wz Aoely, syrisesy
so[ouLIeg Wpooy “eA

Ilse] snop
"JRA “AX 29 “IA Hsepsnop

“AD 29 WG Bessa

"UNUND BURDIXOUI “IBA

OUJIA “IBA
“AIO 2 “YUV oej[lA

"[JB.] SUBX9A

"p,quoo erutsong

54

wejaquin winiuosAjadury

[add], Jo ued] o¢p eiey ‘ZS7EE WOVA se “J sisuoulyo wnuosdAjog "J ‘s}elI] sIsusweATyOUaG
“MOH
[adAy, Jo wed] suNN ‘€Z70LL WOVG ‘ds wnuo3Ajog 2» ‘JaIq esnidsiydure
avaovuo0sAlog |
[adAy, Jo wed] o[epselg “d ein se ‘nN
‘(Ajuo ojoyd 2 apts) €¢979E ANd X9 WOVAG (‘110 ],) sua8und uojAjoepoydaT gjepsejg eurwuiosoA
[adAy, Jo wed] soqpues 3 ‘yaIq Ul
‘(Ajuo oyoyd 2 apl|s) LLpOh UNd X9 WOVA “TJ Wng[n13209 WINIUOUIS]Od “MOH 2p “IC Wuowrsjod
[edd], Jo wed] souor "3 BIWO][OD SB dU99IH
‘(AjUo oyoyd 2 sapl[s) TpI9E ANd X9 WOVA — (YOOH) SIIDeIS Sl1a}SOININ, yoed elequinid
[adAjos]] pZ SUSWIIID ‘pI8h WOVd "yooH wnunieyoind wntuowesog ‘Wy Ul “syorr esojns
[ad Ajos]] "2 PIID se ‘uly
WOVG U! [7p “Ing isuny “zed Y “Gey 2 “YOOH wWnswsosijeo uojAoepoiday ‘wUaH “d ejoony3
[edAy, Jo wed] ssouyieyy *) PIID se
‘(Ajuo oyoyd 2p aplys) “O9LZE UNd X*° WOVA qudaIH (“IPUeg) smeIpIO snyUeUTT "yleH ovis
[adAjosq] CZES “MIOH
‘(Ajuo oyoyd 2 eplys) SLLZE UNd X°9 WOVG ‘uog “5D esojnpurys eljaso0T "MJOH esouny
*P.JUOD sEdIEIUOUIIIOg *"p,juos BIlUIdoNg

53

[adAjo,oxy]
JOjABL, 2 sajouure “O9EOT JOPIeD ‘g9PSpI WOVG

[adA4josy] WOVa
ul [7g¢ ‘doing 18unJ “szeg 2 “IU ‘qey

[adAjos]]
Jeays ‘WOV U! 8Z¢ IsuNy “WY ISAM “UID

[ed41] WOVG UI €€1 ‘Pan ‘wy ‘N “yueg

[adAyosy] 145
"Sxq “xeg “OA “YIOS = 78609 = 1968€ WOVd

[adAy jo
ued] CpOl aseq 2 Advil “19xeg ‘g/99p WOVG

[adAy, Jo wed] [¢9 osyD ‘ggzep WOVa

"WS “qf WNtjoyIpes uNaD

aeaoEsoy

‘Od eedasojday empjed

“JT PIOYISSIOIeU
"V SB ‘S[ON elydez ouowsuy

avadENounueYy

T9MOH ("J) eoIqis enuoWy

ysing eyejosour] eruojAeID

Aled (Aeig “y) eziyiesour eruosAeyD
aBad"[Nj10g

SISUSSUPIYI] “IBA
‘J “YOOH wmejnundwes wmnuoshjog

*p.juod aevadevuO0sAjOg

QJIAeS stueqin

“Zeq eueisegjan
“‘pAS b199}01

QJIABS SENUOU “IBA
"JUD WuOs]IM-a1TeW
"WUIND P[OOTTUO}ARID

‘Wy eyuse

"WUIND eURMOYIIOMy

"p.quoo ejuIDoNg

56

[adAy,
jo ued] WOV U! OS¢I “PAN ‘Wy 'N “queg

[adAjos]] WOVA UW

EQ] “Sxq ‘SsOY Isun J ‘*ZUeI], 2? “Woy *°Z5ef

[adA4juAs] yously “D ‘679LL WOVd

[edAje1eg] Sp9 “MIOH ‘769801 WOVd

[edAjosy] €707 “MIOH “PepLL WOVG

[adAjosq] ZLL1 “AIOH ‘LIPLL WOVG

‘|3nog (Ysing) elOyIpuess eUNtT]a]

“J epnu ea

IBIIUSBAJIXES

"qq septosodoAul uoula\solg

avaoujNy

‘ds wintyey

‘ds wintyey

"19g eljepuod winjasdA90990

avaovlqny

(eueISULIOQgONSNe “IBA

aevloyoney *q) euerIsuliaq

-onsne ‘dss afAes
(a]taes) euelsuliaqonsne

(vonneise
“IVA BeJOYONIY ‘d=)
“pAsS (‘WOy) BoeIse

‘do stuowlajsoe

“MIOH 2 “YY PIX

“MOH
w ‘syorr we[NUINIUOD

“MOH
w ‘syoer tjasdh909909

*p,juos erutsong

57

[18a Jo adAjosy] 29e'T ‘gE8py WOVA

["JBA Jo
adAj00H]] JopleD 2 Epp UsU0XANY “O/spyl WOVA
[sea Jo adAjosy] €7-79 “wuwND ‘O98h1Z WOVd

HH
xo [rea Jo adAjojoHy] eyNseN ‘609EE WOVAG

[sea Jo adA}oFOH] PLOS Aes “L61Eh1 WOVG

[rea Jo adAjo[0OFq]
Jap[eD 2 979 UsuOXANY “pL8ppy1 WOVA

[adAjos]]
O9ZET ayn ‘(AUO apts) CO7LIT WOVG

[jeliajyeur odAy,
xa] JoAeW] 29 JOYONID ‘(ATUO apts) 19p9E WOVA

[‘dss Jo adAjojox] *j vYNSeIY ‘ET9EE WOVA

YUOmMespy ey~pURIOIW eseIyIxeS

“T epnu eyo

“LIOL, susdsaqnd eloyoNe}

‘ssiog eotuodef eseyixes
TOUWI]O}

"IBA “DS ® ‘] lowjo} eseaixes

OPTARS 2 JopyeD Husjny
‘dss ‘[3uq WeA] eseyixes

“UNXEYL JIeT]IXe wntusdsosAIyD
"ITV eIOYIg eselyixes

"WIXeY BOSTY BSBIJIXES

*p,jUOd aBadEseIFIXES

(‘u-"s *g) aftaes
(Aejoseg) sewjueIoIU
-OUSRIJIXES “IBA

QIIABS JOUTU “IBA

SIAC SI[BUOIPIIOW “IBA

ayiaes eoruodef “rea

OTAeS
ILLIISISNJIP “IBA

sd LEN
PUPIO][IPIOD “IBA
‘JaIG (MYOS) seJoyONoY

QJIAeS oeleYy

lokep 2 JYONID Wosyosy

oTAes
~ wnseseyixes ‘dss

“p.quod BruTIoNg

58

[adAy Jo wed) ‘jy ‘swe ‘pyOCe WOVA

[adAy] ounweuy ‘1¢78L WOVd

[ea
Jo adAJ] 9COI “sxq “xeg “OA “WUIND 2p “YIOS

["IeA
jo adAjopOFY] JOPIeD 2% L8p “AMM ‘616rph~1 WOVA

[rea
jo ad4y] WOV Ul 6€ST ‘PAIN ‘Wy 'N “queg

["Ie@A
jo odA4T] WOVA U ZZ Hsun “WY ISAM “WUD

[ ‘rea Jo adAjopoH]] BBL attAes ‘p7SE7 WOVA
["1ea Jo adAjosy]
WOVG U! ZZS1 ‘pan ‘PAS ‘Zrr9e WOVA
[Iva Jo
adAofoHy] JO[AR]L, 2 WHISEZ JOpled ‘1766S WOVA

“sslog esSojAIs eT]AIIA

“Qnisis
useoiAode] “JeA “J Wines eseIIXes

*‘pussoy (Jodig) ejeyodousjs
"IeA Jadig epeyodoineys eseljixes
“JT BNUIID PSLIJIXES

"Tl 3° “Y Sljeuornuajdas

"IWA *[3NOG edIIpuT[AD eseIpIXxes

‘one epidsiy eselixes

"J eyoytsoddo eselyixes
"T BIEN BSeIIXES

WeyeIH PIUISNIII eSVIJIXES

*p,JUOD BBIIVSBIJIXES

‘J *S}PIIF{ SISUSTIONO)

QpIAvg 193-o"seIjIxes

QIALS JE][STUW “IEA

QIABS JOISUOT “IBA

JIIABS DOTYOO]OY “IRA

Qyaeg wniesoyonoy “IBA
"YOOTYOS oeseIpIxes

otAes
eioynisoddo ‘1eA

(Isayny “q) sT!ABS
(‘pAS) Wainy ‘eA

aplavg eoulsntiay
"IBA “JIC, atyYoszed

°p,juod BIUIDINg

59

["1vA Jo adAjos]] OPLOp esoulaoes ‘dss *yooH

UNd X9 prsz ssieg ‘(AJUO apis) ZERSII WOVG X9 ‘[SNoq esOWIdRI SLIv|NIpsg OJIABS BSOWIDOPI “IEA
["1eA Jo "yOoH 2 ‘Wueg QJIAeS
adAofoH] “1B 39 99€07 JOPIeD ‘LE~Ss7Z WOW BOUAYIOYNUIO SIIeNIIpag SeOUAYIOYNUIO “IBA
[‘1ea Jo adAjos]] (Opl “sxq “xes “OAW ELINA
“wUIND 27 “YIOS =) TEp-9S “WUIND ‘¢¢Z71I9 WOVA ‘ZIDY ROIPUL[UIOIS sLIejNoIpag aeoipue[usol3 “IBA
[adAjosq] CELOr
Wd xe wuowryD “(AJuo apis) LLESTI WOVd VT Sisuspeued SiTe[NoIped MUOJUT[D “IBA
[sadAjos]
3 adAjojoHT] (ITpZI “Sxq “xes “oAP ‘wun
R “UlOS =) JOpleDd 77 906 “ANY ‘LOSPIT WOVA “Wuog eS09}0e1q SIIe[NoIpsad OPTARS BSOD}DVIG “IRA
[18a Jo adAjosq] ‘uudg eqje ‘dss QJIAeS
L69E UOSKed “6ELOb UNd X? I8ESTI WOVA "YOOH Xe "[8noq esoulsoes siejnoipag SUq[e “IVA YI9_ WUOJUTIS
[sadAjosq] Ccp JJopsyng ‘WOVd
Ul 8ZT “Pen “Wy "N “4yeg = presol WOVG "[8noq ksoJOUIsU sIUO;EYD “MOH 29 “Jol StUOsYys
[adAzojoH] E661 “Ie 39 MOIPNT ‘ZOZ7LII WOVA UleIg NBUTZ, sluejnoiped aTARy SISUSUEIN]G
[dss jo QJIABS SIsuapeosed
ad AjoJoH]] SIAL 2 66ST JOPIRD ‘IIEhy WOVG ABI “VY HWYSISND BOIUOIOA ‘dss ‘use sisuainqye

avaoetiepnydo.19s *p,jUOd BIUTDONg

60

[adAjos]q] p6p AO 7 Plod ‘pS7T8L WOVA

[edAjojoH]] WNnIedTUOIOA
‘d S® WOV Ul ZpIZ “AlUQ, “OA “WoNy

[sadAjosy] CLZ “MIOH “SOL80T Pure TST8L WOVd

[adAjojoH] BSS¢ aT!Aes “97109 WOVA

[dds
jo adAJ] Wf X9 LOPEZ ISP1IOJ ‘C6p~LII WOVA

[adAy}
AI xe Jozousny ‘(AjUo apl|s) p6pL11 WOVG

[dss Jo adAjosq] Jd X29 AoIBWOY “LLLLy1 WOVA

[dss Jo adAy Jo ped]
(AjUO apts) UNd X9 Y676E UOsIapuy “ELLp1 WOVA

[adAy] Socoe
Wd X92 6SL9T Aetpueisg ‘(ATUO oplys) p6ESTI WOVA

[adAy) L67Z9€
UNd X2 Suipooy ‘(ATUO apijs) 67pS1T1 WOVA

['iea Jo adAJ) 75/17 WABI ‘SLESTI WOVG

"yooxH] ey]Aydosoeul eIstinC

“"T Bliode ejOIOpsed
“SH BSOOSIA BIXNOINOUIET

swepy wieqides siensipedg

WUs “AA “AA SIsueuUNA snoseT

"Wi eIpueUWAD se
"TPULS (PIA) JOUTW sHOseT

“uyJaRy vone]s syOseT]

[purrs (“PITLAA) Jour sHosey

“YOsty
2 ‘yYNoD snoydiyje uowssueg

MBS (ABI “W) B|Aydosoru “dss
MRIS (IUIG) SOplouryLyUe ejjaydoyY

"J voneayAS SliepNoipeg

*p,juod seadetIEpnydoi9s

‘Zuluund eussneped

gIAesg sejOJopoed

‘MJOH 2 “WY seposou

OPARS SITBoIoy

gylaes sisueuund ‘dss

ZuUvI], seipueuwAs “dss

SUNS
SISUQDJBYISWIeD ‘dss

QIABS SISUZOSETe
‘dss ‘zuvly, seipueuwA3

ayiaes avlomjuesep

‘WUIND 2 “YY esesjuos

STARS BBdIVBATAS “IA

“p.quoo erujsong

61

[adAjos]]
WOV U! €6p “9S ‘ated “WOIDIY ‘"S9/A

[adAjosy] WOVA UI Zp6 “AUD “OA “WON,

[adAjos]] ¢28 “MIOH ‘SOILL WOVa

[adAjos]] E9¢ “MIOH ‘09E8L WOVA
[‘dss Jo adAj0s]]
OSOT9 ANd xX? 79-c9 “WUIND *78/Lp1 WOVG
[adMofoH] WOV U! Z6p ‘UUay ‘sxq “y ‘Uey
[‘4ea Jo adAjosqy] jamog

“STE9E UNd x2 (AJUO opts) OOPSTI WOVA

[ad4josT] WOV U! L971 ‘PAN “Wy *N “‘Wueg

"boer (""T) eqopiniwas enayumnisy

IvIIeNLL

“ASO, BOULULIOS BLIBOTIAWW

dVIIVIICWIL |

“7 wnurjuow umnuerjos

SII SIsUdINS eWOIYIO]INIOg

aBvaovULlOS
“QpAy (‘S[ON) sisuosuIWIOAM eAassag
""T BI[OJIZUO] VOTUOIOA,

“YOOH

Xo “WUog eJOYIsuap sisejnoipag

Avly “YY Beqieqiulas slIejNsIpeg

*p,jUOd aevadeTIejNnYydo.1Dg

“MJOH 2 JIG seyaWNIN

SSOA BlusWaNYy)

“MOH
w? “syoes sewoIJYydOoTIN900d

“MOH 2 “Syoer eorne

gjlaesg euerjuourxes ‘dss
“MIOH 2 JoIQ oelUaziNK

OpARS
IEI[OJISUO]-IEIIUOIIA

QIABG BIO[ISUDp “JeA

SUdDSIJNI “IBA
"MOH 2 J2Iq susdsayny

*p,juod eviulong

62

[dss Jo
adAjojoH] WOV UI CLST “PAIN “WY ‘N ““4yueg

[adAjojoH] “u's “MOH
“SSOI “PAN ‘Wy ‘N “Yueg x9 ‘pecLpl WOVd

[adAjosy] 681
“yeD “anulsunq “u's “JOH ‘p8es0l WOVG

[edAje1eg] 7061
‘idag € “O'd “J91DeID ‘WOVC UI “U's “MJOH

[edAjos]] pOr “MIOH ‘L978L WOVd
[adAjosy] 6IZLI ‘MIOH ‘7P08L WOVAd

[adAjos]] G6ILE “MIOH
‘OIE “PAN ‘Wy 'N “Yeg = ZpplLL WOVG

[sdAjos]] 9621 “MIOH ‘9888L WOVG

"JT sisnjed eforA

"MON PTIQEIS LOLA

“wuag eeqQO] E[OLA

*ZJOH] X9 JaXaey BILINIIGIO PIOIA

IBIIEIOIA

"weYyD % “[yOS ejeydasorAw eiddiy]

‘ds eiddry

‘bug 1ojsutid eddy

dEIICUIGIIA

“J suopueds euelIaeA

IVIIVULLIIIVA

QPIAeS eUuLdLIOWe

‘dss “dq (‘wnyds) se[o1A

aplaeg lates

“MIOH 2 “JIC esnjyje

‘yy sisuapeurs

"MOH 2 “WY euseulod

“MJOH 2 “Syoes oe1ew

“MJOH 2 “32Iq BIOUNfuOD

"MOH 2 “SHOU C[[NUIA

*p,juod elUIDONg

63

[edAy, Jo wed]
bynsjelH NIeYORN “17667 WOVG

[adAjosq] °J “sve ‘18667 WOVA

[adAjos]] epeweyx ‘9/667 WOVG

[adAjosq] oursny ‘96667 WOVG

[adAjosq] eynsyenpy nieyoen ‘po0zTE WOVG

[adAjosq] 1ynsey ‘8667 WOVG

"J snseia9 snunig

avaIEsoy

“OONZ 2 “Qalg siuejoned vasueipAY

avaovasueIpAy

oun] ejeUdID sNsey

avoousey

“IONZ A “Qalg SIAJOUTIE BIIIID

aeaIBIY}IID

"lq Wrjeoiny wnuNg!A

avadeTosiide

“IONZ A “Qa sAIOUIJNI IOV

9B998.199Y

(‘o-"d erosdoyay) Isi¢s
(‘s}¥JI) Ises90-opnasd

‘J “SIRI sIpliejonad
-ovosueipAy

epewlex 13ej

"J2Iq louesny

“syel] WnueoqgeAU

‘J ‘SJeIF] asuouRsoyYy

WneLSVINIDONd

64

[adAjosq] Ses “MIOH “6706L = LSL801 WOVG

[adAjos]]
Z£66S UNd XP LS-79 “WUIND “IZI9TT WOVd

[adAyereg] Z9L “MICH “9SL801 WOVG

[edAjos}] WOVC
ZB8I “Pan “weg = €79p BUT ‘1h06L WOVA

[adAjos]] peyT “MIOH “1€06L WOVAG

[adAjosq] eseurysox “E607E NOVA

[adAy, jo wed]
9681 ‘Ides ZZ ‘eyNseII NIVYORN “L706L WOVA

|, BadeSOUL,,

QSOY CURIPIETIM BIOvOY
‘ds eipuei[ed

enbiisnel “T
se ‘Ijueg sisuoweyeq BUIOTISA'T

avaonqey

“ypuag WINIOYIXe] WNuUe|OS

aEadVULLOS

“UNXRY] BI]JoweRdopnasd eNJEMaIs

svoobIt |

‘JON 2 “Gals elssego xeIAIS

aVsIVIAJS

“MOH 79 “WV surisip

‘yxeg HsuTWUINS

"MOH 29 “YY eyeu0zIq

Zuo] eye[nuue

VITAUNGAVA

“syoer IAeMOY

VuISOINIDONd

"J “syerP] reseurysoA

“syelIP, WNuIdeIA}s

*p,juod UNIYSBIUIDIN

65

[adAjos]] 6P8 “MIOH ‘POI6L WOVA

[adAjojoFy] Woy 2 OZL 2]oULIeg ‘01616 NOVA

[ad4y] 4ry “€606L WOVA
[adAjosq] ZpT “MIOH ‘0606L WOVG
[adAjos]] ZEOT “MIOH ‘9806L WOVG
LO6I “E€vI-Zhl (OL Wd

"WY ‘N 99S “YY Sliensai “YY Jo WAUOWOY Jae]

[adAjos]] (11ze1g) OTyT “MIOH ‘0L06L WOVA
[adA4jos]] CSOT “MIOH ‘0906L WOVG
[adAjos]] 1601 ““IOH ‘9S06L WOVa

[adAjosy] WOVC U 6L1 ‘PAIN ‘Wy “N ‘“yueg

[rea Jo adAjosy] 967 ‘MIOH ‘1S06L WOVG

‘ds winiprurmoy

avaoepuldes

yousopy SieuOZIIOY sniodiune

avaoessaidns

, Yonelg-uasousuin3a’T,,
‘IWUsg sIsuso[ndese eWwoOyTIsSAT

‘yuag we[I01Iped eIovoy

“ds elovoy
‘ds vlovoy
‘ds vissea

“SJB eURTINbs} eIOvOy

YOSIOPY SIIoes3 epuenyes

*p,jU0d avaoEqey

“yy TAemypoy
VALS

aojaueg sisusoniqg

VITALSHOU

‘pAS evueiuasioy)

“MOH 2 “Uy sisuajojos

“MIOH A ‘syore ees
“MTOH

wp “syoer sliepnso

“MOH A ‘Syore eouopi

“MIOH w “syoer wyI0RJ

“MIOH 29 “J01q osuedxo

(edA}09 *y) “wun (“MOH

2p ‘yy) edAj09 “IBA
“JIC 2% “YloseT eyeuryso

*p,juod BaUaAtYy

66

[adAje1eg] WOVC Ul ZZ “Sx "10xa

‘a “PAS = TT61 AON 072 ‘BPH “MH *L996L WOVG

[adAjosq]}
WOVG U! 06L1 ‘Pen “PAS = 1996L WOVG

[adAjereg] Cops Joye ‘7E7P8 = 87PS WOVA

[adAjosy] 1681 3dag ‘soson
pue inowlXag ‘eueimopiey [ay “L90L WOVd

[adAjosq] 768

‘dking sading ‘Japsan 2 “sno “O188s WOVd

[adAjosq] 907 UNYsiI9g ‘1E06€ WOVA

[edAj0s]] 069 “MIOH *09S9L WOVG

uyny (“7 wnurinbe winipiiard

wINJOSOJIAD “YW SB
oy (“T) eurwaoj-xipoy UINLIAIY

‘WS “A'f (TP) BOUISIA eIPIEMPOOAA

"J SI]IqIsuas B2]90UO

soyAydopt.13}g

“JT saplonoyey) wnsAdosy

avaounounuey
“PITA esusouIns wINITeIG

dvIIe ge

uaka JOJOIISIOAZ BISSeD

avaonqey

jjney euerowey

“pAg mpueydos

iney Wanque

“"pAS eursiiowe
SISHCONIGHAN
(1 wnrusesydisy)
"ZUR, (JOPSON

2 Sno) WAIdos!

SISCQINDVUHdIaL

‘pAS WMUTIZA9
WNINDVaHdOwavVHdSs
(‘Y PUIOUTA)) “SIeIIH “A
ap “uN (“syoes) WARMjOY

aAVALVUOS

67

[adAjosq] cgeg IEW ‘60L6L WOVG

[adAjosq] c1gz souos ‘Ezoog WOVA

[edAjosy] 669¢ sue ‘OSLEy WOVA

[ad Aj}0s]]
WOVd U! 0611 “[8S “1e1 “WOISTP ‘S94,

[HH x9 adAjos]] towey ‘Ss ‘op/67 WOVd
[ad4jodoy] 19WI0I$ ‘10,6, WOVd

[edAjereg] [Ese JoxIEq ‘SE7PE WOVA

osueyW Wntjuedin} euljnoiedo

IBIDE|NAJOAUOZ)

ezjuny (“sje A\)
sI]eUSPINIO vay]OIUaTTY

avadeIpodousy7)

‘S20] (“M@H) epunquoy eoepiqeiy

avaIvIUOUsIg

“AIQ) Suadies elAnesseN

dBIISTINJA/ICIICIIISY

‘Jeq SoploysnsdAjod eispooM
“PIEAA BotueULES stajdonyNNs
[Sold

C PIA) WneIsnsue wNIAWIY

*p.juod sajAqdopiieg

*pAg eurjnoiodo

"yy sipAyorisomds

suleyy b)99}0p

‘JSOA OPIANESSeU

oqaan

loWey IeISpOOM

JOWIOIS sIpliajdorpnyys

[ney eyeuoIONUTTZUO]

*p,juod stsdourpei,

68

[dss Jo adAjos]]
EL6PE UNd Xe (AJUO aptis) 9E8¢8 WOVA

[adAjos]] ¢c-€9 UosIaled “prSpIl WOVd

[adAjosy] 6p “MIOH “OLL801 WOVG

[adAjosy {] ‘uund ‘p17Z9 WOVA

[ad Ajos]]

E681 Avy ‘urs o[epseig “TEss0l WOVd

[adAjosq] 0¢-99 UOsIaIag “ZpSpIT WOVA

"Mg (“D) dATBAIg UNpIOISOMION
aBoel]V
"J sapueuow wntusjdsy

sayAydopiis}g

"UINO,] IZAMULCY SONSNIZ

avaov0g

‘JS1O,J XBUd) WINIWLIOUd

avaoell’]

‘yuag euldAyeo ajaoeyds

sBoIBIUIvT]

‘pnajg euvoddep sniodiuns

avaoessoidnd

(‘u SddAWOI() BITABS
(‘pAS) Ipsoosoyjou
‘dss ‘Sadg sipuoaeunid

SHOAWONN

‘sIajog vuerYyIOIZe

“MOH 29 “UNV siisnez

‘uuns iw0yd

“MOH
w ‘JIG ejooaoeyds

"slajag eooysode

*Pp,jUOd Opein

69

[adAjosq] °y ‘swe ‘SZTIEE WOVA

[adAjos]]
9[3unld ‘WOVC Ul 8ZZ ‘Hed ‘1a = 6876L WOVA

[adAjosq] EgT¢ “MIOH ‘871601 WOVA

[adAjos]]
WOVG UW C6LI ‘19D “sd “Ad 9 IA

[adAjos]] 878 “MOH ‘€8S6L WOVG
[adAj30sq] [ZI “MIOH ‘8076L WOVG

[pyusyny])
Ie ‘31y “oA ‘99q ‘dads = IZ7ZpS WOVA

‘J ‘YOOH esoused edieoelog

avadelnuedure.)

“MAH Pun[ea eIIOjJoUINOT

IVIdCUISB.IOg

"Jq euepjol o1seuas

IVIUOIIIUIG/9V9I8.19}SY

"YOoH

"yjuog (J) eoorWIesiew sijeydeuy

ava[NU]/aeIIe.19)SY

oye[g{ WiniJopenss wniusWAIEg
Jayeg eAyorjsoyAyd erpidsy
dVIYJUCT IY /IvI98.19}SY

[PUT WiNTAIeIs wWnTy

*p,jUOd aeadeTTV

IRUTYIO]L, A OY sedresesod
“MOH

3 “yIq smuodsoysijop

"YY ejOotuoIdaues

“‘pAg snusoule

"MOH 2 ‘syoes snowaeyds

"MOH 2 ‘syoer oerpidse

sijiaaeunid ‘dss

*p,jUOd SddATIOIN,

70

[sea Jo
adA}0j0H] WOV U! 96LT ‘PAIN “Wy 'N “Weg

[adAjosy] [S71 “MIOH ‘S7TS6L WOVA

[snuetny]
QI ‘sry “OA *90q ‘'3adg = 80LPS WOVG

[edAjosy] Zepl “MICH *7616L WOVG

[sodAyereg] ‘yoodso1 CCZ] pue CO6I “9S9T
‘S661 “MIOH ‘9€96L “SE96L ‘PE96L “€£96L WOVA

[adAjodoy] 861 “4IVIW ‘L676L WOVG

[edAjosq] WOVG U 76 “doing 13unJ “qey

Dg
(Ady) snousosiyes snditog

avaoei0dfZa
‘u30Z eBIeUID) eIUOdeARD

"Iq eoOsIoy seutsodsouelsy
"U30D eURISUIUIEM eIINsUy

avaouvIqanons

ea0wody

JVIIE|NATOAUOZ)

“bIW
X9 ‘Jan, Wnipuejued wnwsUdATOg

avaourpodousys

"J suejnu sUgTIS

seaoeAydoAleD

QPIABS SNIIUIOJI]ED
"ea ‘dad snuevdiiowe

“MJOH 2 “SHOU smeI

*Badg SNUIISsIAOU

“MJOH 2 “SyoeL oelinsue

“MOH 2 “SYOVL SNUIOIA

“dt Woy tursusAjod

yose’] snjyeinboeul

*p,juod sadAUI01—)

dad

{edAjodo] snjejjoinos
‘Nl S® WOVG UI 6901 ‘19D “dA “40 “IG

[adAjosy] 1/781 “10H “6656L WOVG
[adAjosy] 069€ “MIOH “STI601 WOVG

[adAjosy] €ZEE FIN ‘67P6L WOVA
[pnuayny] O88I ‘soraUSID “f

‘LI ‘81y “oA “99d ‘*8adS = LOLS WOVA

[adAjosq] €601 “MICH ‘P916L WOVG

[adAjos]] ETT “MIOH ‘pSp6L WOVG

[‘dss
jo adAjosy] WOVA U! LL07 ‘doing 1sun,j ‘“qry

‘wyasuq euRjUOW eIqioydng

‘ds Jomlueyy

“TJ suain eydoner
II sesuNes JomULPY
“q BlIBDOROX|”

Se ‘]Janyy esojnpuerysiq unides

‘TEN JO[OOUOD UWOUI9}SOUTDY

avaoviqioydny

yuny veexa erodsoyouAyy

“7 snumjisew sndiiog

*p,juod aveovsedA7

*pAS peyoszuedy

“MOH
aw? “syors snpuese]o}

*MJOH 2 “J9Iq snuvsexeo

"UUs
*q oesurjeo-smoyuew

*Sadg snurosousid

“MOH
zw “syour sIUOW}soUTIIe

“MIO 2 “syoes snour}d9/q0
ayIAeg snoyoseou
‘dss *J90I1Y9S

(‘WSdq) sneie[OoUT]

*p,jUOd SdATIOI(

72

[adAje1eg] Ooxayy ‘eoeAeUIOND
38 9681 “Id9g ZZ ““MIOH “E1601 WOVG

[adAjos]] WOVC U! Z6€7 “PAIN “Wy °N “Weg

[sea Jo adAjofoH] 7987 aT!AeS “1000r1 WOVd

[adAj0sy] WOVA UI 9p Bun “Wy *N ‘SHIA
[oueyyny] uosuIqoy ‘1766 NOVA
[adAjereg] 6LIE “ICH ‘€60601 NOVA
[adAjo0sq] gg0¢ “MICH ‘760601 NOVA

[adAyudg] CELI “MIOH “PEZ6L
WOVd pure [adAjuAg] [081 “MIOH ‘S€76L WOVG

[adAj}0s]] 090S ““IOH ‘6L0601 WOVG
[adAjosy] wes ‘HH X9 9967€ WOVA

[adAjos]] WOVA
Ul 8SI-LST ‘Sxq ‘ssoy ‘gy “*ZuRIy ‘Woy ‘ORL

‘ds winipoulsoq

a CD Bury ezopedzey

Aqouleg (‘U19.J) SBAOU-9e119}
"IRA “OQ (‘7 sisodues sidonAxO

‘ds wintpowsaq

‘yluog eyj|Aydououwl eIsJoquopleH
"SAH eyoyoind eruesojod

"HJ PULIXOW BIIONI[D

azjuny (‘WUsg) susdseqnd *g pue
azjuny (“J) euRIUIsIA BAINGpelg

"syeqy to[sutid eruryneg

IYSIA Slepnsue snjosseyg

‘INUdg SIsuINWe SIPNseIpEpy

avo0eqey

“MIOH 29 “2Iq snuedxoul

(‘d-"| erutsong)
"ysey (“MY9S)
styusquinsod-sezapedzo}

aytaes stdonAxo
“Jea ‘ylasey] snowuoddey]

(d-"y

eiursong) “ey (-A#YdS)
neynowued-iesApoy
‘di yow-w ovissoquopsey
‘yy oviuesojoo

"yy 2eLION]9

“MJOH 29 ‘syoes oeAIngpesq

‘wy ejoonuyneq

BLIP, Ҥ Bjoorynze

“"WOy SISUSINWe

*p,juod sadAWI0.172)

73

[adAjos]] “}UOP WyoUAIAsSIS
" S®@ WOVG U LETe Wung ‘wry “N ‘Aq 9 “II

[edAjosq] WOV U! 8207 ‘doing 13uny ‘-qey

[adAjosy] E711 “MIOH “OLP6L WOVG

[adAjosq] g9¢ “MIOH ‘pes6L WOVG

[adAjosq] eseurysox ‘6666, WOVd

[adAjosy] Z7S1Z “Tsesg “QI9H ‘21M “SEs6L WOVA

[adAjosq]
arepseig ‘WOVC Ul 96 ‘Pain “Wy “N “yueg

(‘3 wintuds[Q =) ‘[3sn0g
wnoyIpuels wmniyouAyAsisg

"THA Snoyig sno0j1Z

svaoEpli]

"MEP PYOJIpuLI. elysolog

evan elsoddiy

‘d BY wMrpoyiqye saqry

dBIdELILINSSO.1I

"ye PURTYOYLYS syseipelD

“uog (“[solq) elojtuefes elio[D

"esy “¢ snipoyney, snuidny]

*‘p,juod avaoeqey

‘yy snqoid

"sstq 19019

“MIOH “Syore oetysosod

“MOH 2 “SyoeL eJOIIQ

ouessny snueryoyrys

(‘Ja1q Idered0INoU
"AQ = ) UUs ‘gd smeNsol

"JIC. sI[eIUepI990

*p,juod sadAu01)

74

[adAjosy] WOVA U! ‘pais[eH smensip
"A. se L7ZZ ung ‘wy *N “AT “1

[adAjosy] SpE uosyors ‘(€IZ6L =) EPIET WOVG

[adAyo0H]] Ep-9p ONEAe “98981 WOVG

[dss Jo adAjojoH]] UOIMEN “L79¢ WOVG

[‘dss Jo adA}ojoH] Lpp aAes “OSpewT WOVA

"AMAL payeusisep Cp¢ sou nq “COL “S99
‘CRT “MOH epnpoul FOV Ul sedAjered Joo
:310N ‘[adAyereg] 6€S “MIOH “€08801 WOVG

[adAjos]] 8cel “MICH “0776L WOVG

ZUCMS (“]) BUIBIIA PISIOIT
SOW (“TT StWIOJSvONID BIUULUAIIG

avaoeog

"7 BURIUTTOIeS WNIUOWTT

“PIAA CIA eumiieu eLoully

“PIAA CIINAD PwHeUl eLIouIy

seqoeuIsEquIN|d

‘ds snyjuelo7y

‘ds snyjuelo’y

avadeyjUeIO’y

“L 9d Hparsjey

“syoer avluURUyI9qQ

SIQUUOD 2 IAPS
TUBIUT[OIVS-UOUT|

sIouu0d
2 alaes snoyioed “dss

s1guU0d
2 a[IAeg eoluospny ‘dss
"AX ([YOS) selowise

“MIOH 29 “UV snisos

“MIOH 2 “SARL TYULIO]

*p,juod sedAU01—)

75

[edA J] 499g ‘SAN X9 (ATUO OpI[s) OpS9p WOVG

[‘1eA Jo adAjosy]
WOVG U! Z€ZZ Isunj “Wy *N “Aq 7 “II

[Iva JO adAjooFy]
WOVG U T6S¢ ‘19D ‘A “Teyleg 29 “YUeg

[eA Jo
adAyopoHy] IPNeIS 2 ZZp JO[ARL “OIEOII WOVA

["Iea
Jo adAjojoHy] siouuoD 2 sures ‘90r1 WOVA

[adAjos]] naoMey ‘EOOPI UNd X9 S6ZOIT WOVd

[edAjosq] EOI] [eduey ‘cov6L WOVA
[dss Jo adAjojoqy] OSTS a1AeS ‘68EISI WOVa

[adAjos]]
MOLeY “WOV U Obz sung ‘wy *N “II

ysing eyeosour] eIUO)ARTD

dvIIBIEIN}IOg

“HOL WNye]Iquin wWNuOsOIIg

‘YNN WndooIoW WNuUOsZOIIg

‘[3noq wnealu uNuOsOIIg

‘WNN Wary wnuoso1ug
‘WS “Af Snioyiased wnuoso1g

avaIvUOsAlOg

“7 g[euinsues wnoueg
ausaIH (J) wnyjeoids sijyonsig

*s wnJAdozig se
auaaID (“J) wingesids sijyonsiq

*p,jU0d a¥vadv0g

49d 2 “WD seIU0)ARID

QPIARS HeTJoquin “eA
o]TAeS

snsolodiinjd *1ea
OPIABS BOAIU “IBA

QJIAeS JOfeul “eA
“OYOD smedur

apaeg 1Aqsiq

josuey siyeuinsues-io1ued

QJIAeg esojnsni “dss

"[Je snueryood

*p,juo0d sad{u01p,

76

[adAjos]] $g¢ “MIOH ‘7PS6L WOVA

[[eliojyeu
ad4q) S80€ ‘Ski “QI9H X9 21N ‘91S6L WOVA

[adAjosy] eseurysox ‘9676L WOVA

[adAje1eg] OOpT UOsUIqoY “L0L6L WOVG

[‘n saokwio1 jo adAy] sadapurig

‘6L6PE UNd Xe (Ajuo sapifs) pos9p WOVA
[dss

jo adAZ] WOVA U! 60CE “INF “A UI“ 9RN
[dss jo

ad4Z] SOLZI Aces ‘SEL9p UNd X? 9LS9v WOVA

[‘dss Jo adAy]
Aasjay ‘WOVC YI ELEl “10D ‘Ad “Ad “TA

‘ds eoryoidyes

svIIVULLOS

ein e1noysAsg

“J oulede winiyey

“bil X9 [Tany eIpuesijo ejniedsy

avaoRIqny

uOsUIgoYy
({J2MOP) BURIQUIN[OS eISIMIT

“1IO], BIeTJoquin vanseids

“IO], He|Jaquin evanseids

Ysing BAIAQPal PISIMIT

*p,JUOd sBId"d"INII0g

"MOH
aw ‘syors seoryoidyes

‘uuaH] ae1oysAsd

“yaiq mes

‘dpyow oe[niodse

smiun ‘dss

OTARS

(‘yze}) eenselds ‘dss
gyaeg snoyioed ‘dss

aylaeg sisuouejuoUl
‘dss yoog smun

*p,juos sodAUl01Q,

77

[adAjosq] jowueg ‘1 p.761 WOVG

[adAje1eg] 390 “MIOH ‘6Z71I9L WOVG

[adAjosy] 7830S “MIOH ‘9€1601 WOVG

[adAy, Jo wed] ‘J ‘yey ‘SOLZTE WOVA

[Spuayny] LO61 “JEW 81 ‘Yous, ‘21/62 WOVG
[adAjosq] uosuiqoy ‘71462 WOVG

[dss Jo adAJ] gp6l “JeW
p@ ‘Twemsesuey 2 ueuysiyewry ‘TE9¢p WOVA

‘ds uowisjs]e9g

“SIGH soproydiowe enpreyusshq

"SIBAA BSOJOqns esAydiq

aveoeqegy

"USOD BIBUOIONU BLIIOTIW

avaIeRVIGININZ

‘IWUSg euRUOUT, BIDROy

"XT eieqieap eiovoy

avaoeqey

"J snusejes sniooy

avaouluesieds

(‘d eruisong) *
39q (‘[se4) stuowias|ejod

(‘Yy elodsoryed)
"YY (yy) Aemjoy

(‘p elodsoryea)
“wuUND (yy) sesAydip

(- erutoong)
Aeiny 2 0 (‘ysey 39
‘J “SJBINE) stsuouesie

SIXAdOUN

‘dryow xeduns
‘dr yoWw winsodsiq
WhidWIOAWOoUNn

optAes
3? sajouleg snoneise ‘dss
49d 3 “IID Wuedseds

*p,ju0d sadAMI0I

ea, Mie
‘es ds
pat hy {

MYCOTAXON

Volume LX, pp. 79-102 October-December 1996

SOME STEMONITTALES
C(MYXOMYCETES) FROM THE
STATE OF TLAXCALA,
MEX TCO

M. RODRIGUEZ-PALMA
and
A. ESTRADA-TORRES

Laboratorio de Micologia
Centro de Investigacién en Ciencias Bioldégicas
Universidad Autdénoma de Tlaxcala
(Postal Address: Apdo. Postal 63-389.
México 02800, D.F., México).

ABSTRACT

Twenty species and two varieties of Stemonitales were
found in six localities from the State of Tlaxcala, Mexico.
Schenella simplex Macbr., Diacheopsis insessa (G. Lister)
Ing, and Stemonitis microsperma Ing are recorded for the
first time for the Neotropics, and S$. hyperopta Meylan and
Comatricha rubens A. Lister are new records for Mexico. The
collection of Schenella simplex is the first made outside
the type locality in Yosemite National Park, California,
U.S.A.

RESUMEN

Se estudiaron veinte especies y dos variedades de
Stemonitales procedentes de seis localidades del estado de
Tlaxcala, México. Schenella simplex Macbr., Diacheopsis
insessa (G. Lister) Ing y Stemonitis microsperma Ing se
registran por primera vez de la zona Neotropical, y S.
hyperopta (Meylan) Nann.-Brem. y Comatricha rubens A. Lister
son nuevos registros para la Republica Mexicana. La
coleccién de Schenella simplex es la primera realizada fuera
de la localidad tipo en el Parque Nacional de Yosemite,
California, Estados Unidos.

80

INTRODUCTION

Most of the studies on Myxomycetes from
Mexico include a list of species based on a lack
of regular sampling in time and space. For this
reason, the species composition of Mexican
myxomycete communities is only partially known.
The best explored areas are found in the States of
Veracruz, Jalisco and Nuevo Leon (Braun & Keller,
1986; Trujillo, 1988; Trujillo et al., 1986; Ogata
et al., 1994), but only a few records have been
made in other areas. In the State of Tlaxcala,
Central Mexico, there are sixteen species of
Myxomycetes recorded in the literature (Braun &
Keller, 1976; Keller & Braun, 1977; Villarreal,
1990; Hernandez-Cuevas et al., 1991; Galindo,
1992), none of which belong to the Stemonitales.

In this study, we list and discuss the
species of the Order Stemonitales which we have
found since 1988 in the State of Tlaxcala.

MATERIALS AND METHODS

Most of the specimens studied (98%) were
collected between 1988 and 1991, from Abies-Pinus
forest, in "Canada Grande", Malintzin Volcano,
State of Tlaxcala, Mexico. The remainder of the
specimens are from five other localities in this
State.

Delimitation of species follows Martin &
Alexopoulos (1969), Farr (1976), Kowalski (1970;
1975), Lakhanpal & Mukerji (1981), Bersan (1990)
and Nannenga~Bremekamp (1991). In some cases, the
colors of the structures in the descriptions were
based on the Munsell color chart (Munsell, 1992).
The new records for Mexico and some unusual
species are fully described; others are only
listed or discussed when deemed necessary.

Specimens are deposited in the Mycological
Collection of the Herbarium "TLXM", Centro de
Investigacién en Ciencias Biolégicas, Universidad
Auténoma de Tlaxcala, with some duplicates at the
ENCB (Escuela Nacional de Ciencias Bioldégicas,
Instituto Politécnico Nacional. México, D.F.) and
IZTA (Escuela Nacional de Estudios Profesionales,

81

Universidad Nacional Auté6noma de México.
Tlalnepantla, Estado de México) Herbaria.

The following abbreviations are used for the
names of collectors : Estrada Torres (ET), Galindo
Flores (GF), Hernandez Cuevas (HC), Montoya
Esquivel (ME) and Rodriguez Palma (RP).

DESCRIPTION OF THE MAIN STUDY AREA

The State of Tlaxcala is found in Central
Mexico, between 97° 37’ 07’ and 98° 42’ 51°’ West
longitude, and 19° 05’ 43’ and 19° 44’ 07’ North
latitude. The highest elevation in the state is
Malintzin Volcano, which is in the southwestern
portion, and is situated between 97° 55’ 42’ and
98° 08’ 07’ West longitude, and 19° 08/'36’’ and
19° 19’ 98’ North latitude.

A great cliff called "Cafiada Grande" or
"Fractura Central" is found on the east side of
the Volcano. The natural vegetation in this place
is found above 3,100 m and is dominated by an
Abies religiosa forest. In some places. there is
Pinus-Abies forest, with Pinus montezumae and P.
hartwegii mixed with A. religiosa. At 3,250 m or
higher, P. hartwegii grows in pure stands (Acosta
& Kong, 1991). The soils are gravely-sandy
fluvisols with fragments of stones (CETENAL, 1975
A). The climate is temperate between 2,500 and
3,000 m and cold at 3,000 or higher, and the rainy
season occurs in summer (May to November). In the
cold zone, the annual mean temperatures are
between 6 and 12°C. (CETENAL, 1975 B).

LIST AND DESCRIPTION OF THE SPECIES
Family Schenellaceae

Schenella simplex Macbr.

Figs. 2A and 3A.

Pseudoaethalium oval in outline, covered by
a thin continuous membrane, pale gray, sometimes
bright; sporangia cylindrical or columnar;
peridium thick, persistent at the apex and at the
base and forming cap-like structures, evanescent

82

elsewhere; capillitium brown to yellowish brown
when viewed by transmitted light, interwoven,
coiled and forming a columnar helix (Fig. 2A);
spores black in the mass, dark yellowish brown by
transmitted light, globose to ovoid, 5 - 6 pm in
Giameter, with a thick wall and a verrucose
ornamentation, up to 1 wm in height and forming an
irregular reticulum (Fig. 3A), with a conspicuous
germinal pore. Plasmodium unknown.

Habitat: On dead leaves of angiosperm, in
Abies~Pinus forest

Specimen studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 13 June,
1990, 3300 m, RP # 1775.

Observations: Schenella simplex is a rare
myxomycete species and was collected only once
during our two-year study. Unfortunately, the
specimen found in Tlaxcala is badly damaged and
consists only of the remains of one
pseudoaethalium with a few intact sporangia.
Nevertheless, this collection matches the
description given by Martin & Alexopoulos (1969).
As Martin et al. (1983) had previously noted,
Schenella has been frequently considered as a
doubtful myxomycete. Obviously, one fragmented
specimen cannot elucidate the status of this
genus.

There are two species of Schenella (Martin &
Alexopoulos, 1969) both previously known only from
their type collections. Schenella simplex differs
from S. microspora Mart. by the color of the
spores which are reddish in the latter species. In
S. microspora, the peridial remains are less well
developed than those of S. simplex, the
capillitium is rugose and the spores are smaller
with less prominent ornamentation.

Family Stemonitaceae

Colloderma roblustum Meylan

Habitat: On decayed wood and bark of
conifers, mosses and liverworts in Abies-Pinus
forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 27
September (Sept.), 1988, 3450 m, HC # 707; 11 January

83

(Jan.), 1990, 3250 m, GF # 1241; 3300 m, GF # 1256; 9
February (Feb.), 1990, 3200 m, GF # 1334; 3250 m, GF # 1335;
3500 m, GF # 1401; GF # 1409; 11 March, 1991, RP # 195].

Diacheopsis insessa (G. Lister) Ing

Figs. 1 and 2B-D .

Subplasmodiocarpous or sporangiate, sessile
(Fig. 2B), 0.4-1.3 mm in length, 0.2-0.4 mm in
height; peridium membranous, translucent, nearly
evanescent, grayish blue or iridescent with gray
or purplish tints; hypothallus membranous,
inconspicuous, hyaline; capillitium sparse and
elastic, grayish to black, formed by branched and
anastomosed threads, arising from the base of the
sporangium; capillitial threads, 1.0-1.5(-2.0)um
in diameter, dark gray by transmitted light, with
free ends and many elliptical to rounded dark
nodes, giving a beaded appearance (Figs. 1 and
2C); nodes (1.5-) 2.5-3.0 (~4.0) pm in diameter;
spores black in the mass, dark gray by transmitted
light, globose, spinose, 18-19 pum in diameter
(Figs. 1 and 2D). Plasmodium unknown.

Habitat: On decayed wood of conifers, in
Abies-Pinus forest.

Specimen studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano. 14
April, 1989, 3500 m, HC # 1025,

Observations: There are twelve species that
belong to the genus Diacheopsis, most of them
known only from the type specimen (Kowalski, 1975;
Mitchell, 1995). For this reason the genus is
poorly understood. Diacheopsis insessa is very
Similar to D. effusa Kow., but the latter species
has an almost smooth capillitium which firmly
adheres to the peridium by membranous
infundibuliform expansions, and smaller spores
(Kowalski, 1975). Diacheopsis nannengae Mor., I1l.
& Heyk. (Moreno et al. 1989) is very close to D.
insessa, but the brown capillitium, together with
the 9-11 pm diameter spores, distinguish the
former species.

Enerthenema papillatum (Pers.) Rost.
Habitat: On decayed wood and bark of
conifers and mosses in Abies-Pinus forest.

Fig. 1. Capillitium and spores of Diacheopsis insessa

Specimens studied: STATE OF TLAXCALA. Ixtenco
Municipality, east slope of the Malintzin Volcano, 27 May,
1988, RP # 35; 8 July, 1988, 3250 m, GF # 114; 3300 m, GF
# 203; 4 August, 1989, 3400 m, RP # 1146.

Stemonitis axifera var. axifera (Bull.)
Macbr.

Habitat: On decayed wood of both conifers and
angiosperms, mosses, lichens and liverworts, in
Abies-Pinus, Pseudotsuga-Abies-Pinus forests and
riparian mixed hardwood forest.

Specimens studied:STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 26 March,
1988, 3350 m, GF # 823; 21 May, 1988, GF # 31; 8 July,
1988, 3250 m, RP # 129; 3300 m, GF # 109; 13 July, 1988,
HC # 150; 3100 m, RP # 148, RP # 150; 3350 m, RP # 240; 14
July, 1988, 3350 m, RP # 191; 30 July, 1988, 3150 m, GF #
215; 12 August, 1988, 3250 m, GF # 339; 3450 m, HC # 366;
1 Sept., 1988, 3250 m, RP # 352; 15 Sept., 1988, 3500 m,
RP # 410, HC # 476, HC # 485; 16 Sept., 1988, 3100 m, RP #
534; 27 Sept., 1988, 3400 m, GF # 653; 4 Nov., 1988, 3150
m, HC # 881; 21 June, 1989, 3300 m, RP # 1070; 17 July,

85

1989, 3100 m, HC # 2079; 21 July, 1989, 3150 m, GF # 1694;
3300 m, ARP # 1120; 4 August, 1989, 3250 m, GF # 1075, GF
# 1734; 18 August, 1989, 3300 m, RP # 1875; 9 Feb., 1990,
3450 m, HC # 1702; 9 March, 1990, RP # 1526, RP # 1543, RP
# 1551, RP # 1554; 23 March, 1990, 3350 m, AC # 1851; 19
May, 1990, 3400 m, HC # 1894. Yauquemehcan Municipality,
Atlihuetzia Falls. 20 July, 1989, RP # 1079. Terrenate
Municipality, Villarreal. 6 Oct., 1989, ME # 572.

Stemonitis axifera var. smithii (Macbr.)
Hagelst.

Habitat: On decayed wood and bark of
conifers, mosses and lichens, in Abies-Pinus, or
Abies-Pinus-Quercus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 8 July,
1988, GF # 69; 14 July, 1988, 3500 m, HC # 156; 12 August,
1988, 3450 m, HC # 365, GF # 389; 2 Sept., 1988, 3300 m, RP
# 401; 15 Sep., 1988, 3500 m, RP # 418; 16 Sept., 1988, 3150
m, GF # 589; 27 Sept., 1988, 3200 m, RP # 641; 3450 m, GF
# 683; 3500 m, RP # 550; 28 Sept., 1988, 3150 m, HC # 779;
21 April, 1989, 3350 m, HC # 1134; 4 August, 1989, 3400 m,
GF # 1741; 31 August, 1989, 3400 m, GF # 1123; 20 Oct.,
1989, 3350 m, HC # 1509; 9 Feb, 1990, 3250 m, GF # 1314;
19 April, 1990, 3250 m, GF # 1492. Terrenate Municipality,
Villarreal, 27 July, 1989, GF # 1726. Tlaxco Municipality,
Rancho Escondido, 8 Sept., 1989, HC # 1464, HC # 1465.

Observations: Martin & Alexopoulos (1969)
considered this variety of S. axifera to be a
separate species. However, Farr (1976) refuted its
specific status, because all of its features,
except the size of spores and sporangia, are
basically the same as those of S. axifera. In the
specimens from the state of Tlaxcala, we find a
great variation in these characteristics,
intergrading from small to large spores and
sporangia, and from very close to open surface-
net. This suggests that all our specimens belong
to the same species. For this reason, we applied
an analysis of variance to the measurements of 50
spores for each studied specimen, finding that the
means of at least some collections were different
(p= 0.05) from the remainder. To test for
significant differences we used the Tukey test. We
find that the specimens listed under S. axifera
var. axifera have spores with mean sizes different

86

from the specimens listed under S. axifera var.
smithii. The specimens with small spores always
have small sporangia; however, in specimens with
medium to large spores, the size of the sporangia
is variable. The result of the statistical
analysis suggests the presence of two independent
taxa but we consider it unjustifiable to segregate
two species on the sole basis of spore-size and
prefer to follow Farr (1976), considering the
specimens with smaller spores as a variety of S.
axifera.

Stemonitis confluens Cooke & Ellis

=Symphytocarpus confluens (Cooke & Ellis)Ing
& Nann.-Brem.

Figs. 2E-F.

Pseudoaethalia 1.5-3.5 mm height, 1-5 cm in
Giameter, deep black; sporangia 0.5-0.8 mm in
diameter, cylindrical, sessile to almost sessile,
fused in the middle by membranous expansions,
separated at the tips and bases; hypothallus
membranous to more or less coriaceous, purple
brown to yellowish; stalk when present short, up
to 0.5 mm, black, irregular; peridium fugacious
except for some rounded plates which are reddish
brown in transmitted light, smooth and
interconnecting the capillitium between two
adjacent sporangia (Fig. 2E); columella black,
glossy, irregular, dissipating before reaching the
top of the sporangium, sometimes branched and then
with membranous expansions between two branches or
completely flattened and perforated; capillitium
an open and irregular meshed reticulum of
flattened, rigid, glossy and black threads which
arise from the columella, dark reddish brown by
transmitted light, with membranous and enlarged
axils; spores black in mass, dark purple brown in
transmitted light, 11-14 pm in diameter,
spinulose, usually free but sometimes forming
groups of 6-16 spores (Fig. 2F).

Habitat: Mainly on bark but also found on
dead wood of conifers and lichens, in Abies-Pinus
forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 30 July,
1988, 3100 m, GF # 211; 1 Sept., 1988, 3100 m, GF # 420;

87

16 Sept., 1988, 3100 m, GF # 594; 19 Dec., 1988, 3100 m,
GF # 854; 7 July, 1989, 3100 m, GF # 1672; 9 Feb., 1990,
3100 m, GF # 1310.

Observations: The Mexican collections fit the
description and figures of Stemonitis confluens
given by Martin & Alexopoulos (1969), except in
the size of the spores, which they cited as 11-12
um. This species was considered as Symphytocarpus
confluens by Nannenga-Bremekamp (1991). Our
specimens fit the description given by Nannenga-
Bremekamp (1991) except in the slightly larger
size of the spores which she reported as 10-13 um.
The Mexican specimens also differ from Nannenga-
Bremekamp’s (1991) description in always having a
well developed columella with enlarged membranous
expansions. Most of our specimens have free
spores but specimens GF-211 and GF-1310 have
clustered spores. Martin & Alexopoulos (1969)
mentioned the existence of a variety of Stemonitis
confluens with grouped and somewhat larger spores
from Japan, but they were not able to decide if
this form was a different species or just a
variant of the same species. In this sense, the
studied collections are the answer to that
problem, due to both variants (free or grouped
spores) were collected always in a small area on
the same decaying log. This suggests that they
represent different generations of pseudoethalia
growing under the influence of different
environmental conditions. Lépez et al. (1981) had
previously reported this species from Mexico, and
they illustrated both types of spores (free and
clustered) apparently from the same specimen,
although they did not otherwise draw attention to
this feature in their description.

Stemonitis fusca Roth

Habitat: On decayed wood of conifers, bark
and mycelium, in Abies-Pinus and Pseudotsuga-
Abies-Pinus forests.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 8 July,
1988, 3200 m, HC # 126; 3350 m, RP # 117; 27 Sept., 1988,
3300 m, GF # 892; 3450 m, RP # 586; 26 Nov., 1988, 3150 m,
HC # 944; 19 Dec., 1988, 3250 m, GF # 860; 4 August, 1989,
3200 m, RP # 1857; 9 Feb., 1990, 3300 m, RP # 1447.

88

Fig. 2. A. Schenella simplex (RP-1775): capillitium (X 44).
B-D. Diacheopsis insessa (HC-1025); 8B. Sporangia (X 22);
C. Capillitium (X 250); D. Spore (X 1100). E-F. Stemonitis
confluens; £. Capillitium showing peridial rounded plates
(p) (GF-594) (X 44); F. Group of spores (GF-1310) (X 440).

89

Terrenate Municipality, Villarreal, 23 Feb., 1990, ET #
2922.

Stemonitis hyperopta Meylan

=Stemonitopsis hyperopta (Meylan) Nann.~-Bren.

Figs. 3B and 4A-B.

Fructifications in small groups, stalked,
cylindrical to ovate-elongate, with obtuse apex,
2.5-6 mm in total height; sporangia 1.5-3 (-4)
mm x 0.5-1 mm, light purplish brown to reddish
brown; hypothallus reddish brown to dark brown,
membranous; stalk black, bright, cylindric, 1/4 -
1/2 of the total height; columella continuous with
the stipe, reaching the apex of the sporangiun,
tapering and dissipating into the capillitial
branches; capillitium loose, sometimes with
primary branches, forming a semi-open net inside
and a thin and delicate net on the surface,
breaking easily and then with many free paler
ends; spores light red (Munsell: 2.5 YR 4/2) to
reddish brown (Munsell: 5YR 3.5/2) in mass, very
pale by transmitted light, globose, with a faint
and incomplete reticulum formed by fine bands,
5.5-8 um in diameter (Figs. 3B and 4A-B).
Plasmodium unknown.

Habitat: Mainly on decayed wood but may also
be found on fungus mycelium, moss leaves and
foliose liverworts in Abies-Pinus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 8 July,
1988, 3250 m, RP # 127; 13 July, 1988, 3200 m, HC # 128;
14 July, 1988, 3500 m, GF # 157; 18 August, 1988, 3250 m,
GF # 1754; 1 Sept., 1988, 3200 m, HC # 415; 16 Sept., 1988,
3250 m, RP # 502; GF # 562; 3500 m, HC # 619; 28 Sept.,
1988, 3100 m, HC # 784, HC # 791; 28 Oct., 1988, 3100 m,
HC # 787; 7 July, 1989, 3100 m, HC # 2014, HC # 2016, HC #
2017, # 2019; 21 July, 1989, 3350 m, RP # 1114; 18 August,
1989, 3500 m, GF # 1773; 31 August, 1989, 3250 m, RP #
1192; 11 Jan., 1990, 3150 m, RP # 1334; 9 Feb., 1990, 3500
m, GF # 1413; 19 May, 1990, 3250 m, GF # 1524; 3400 m, RP
# 1734; 13 Jun., 1990, 3250 m, GF # 1560; 3350 m, AP #
1778. Mariano Arista Municipality, Nanacamilpa, 25 August,
1989 GF # 1096.

Observations: The Mexican specimens have
slightly larger stalks than those reported by Farr
(1976) and Martin & Alexopoulos (1969). The

90

superficial mesh at the apex of the sporangium is
very fragile and breaks easily. Macroscopically
Stemonitis virginiensis Rex resembles Stemonitis
hyperopta but it can be separated by its darker
color, spore ornamentation and hollow stalk.

Stemonitis microsperma Ing

Sporangia stalked, cylindrical-ovate to
cylindrical, up to 1.6 x 0.5 mm; stalk black,
shining, 1/3 of the total height, cylindrical or
tapering gradually to the apex; hypothallus
membranous, dark reddish brown; columella black,
cylindrical or tapering and dissipating into the
apical capillitium; capillitium arising along the
columella as wide threads, branching dichotomously
and forming an open net inside the sporangium, and
close and delicate at the surface; spores purplish
brown in the mass, pale brown by transmitted
light, globose, 5.0 um in diameter, faintly
ornamented with a reticulum of delicate bands.

Habitat: Growing on moss leaves, in Abies-
Pinus forest.

Specimen studied: STATE OF TLAXCALA: Ixtenco
Municipality. east slope of the Malintzin Volcano. 20 Jan.,
1989, 3250 m, GF # 898.

Observations: This species is distinctive
because of the size of its sporangia and its small
faintly banded-reticulate spores. It seems to be
a very infrequent species, often growing on leaf
litter.

Stemonitis nigrescens Rex —

Figs. 3C and 4C-D.

Sporangia crowded, stalked, cylindrical,
rounded or slightly acuminate at the apex (Fig.
4C), 1.3-3.0 mm tall, 0.3-0.5 mm in diameter, very
dark grey (Munsell 5YR3/1 - 7.5YR3/1) to dark
brown (Munsell 7.5YR3/2); hypothallus membranous,
pale brown to reddish brown, continuous under the
tufts. Stalk shiny black, 1.0-1.5 mm, 1/3 - 1/2 of
the total height, hollow; peridium fugacious;
columella apical, tapered and dissipating at the
apex; capillitium arising from all the columella,
formed by thin threads, dark brown to purplish
brown in transmitted light, anastomosed, forming

91

an intricate net; surface net delicate and
sometimes breaking easily; spores dark reddish
brown in mass, dull brown in transmitted light,
globose, 9-11 wm in diameter, with an angular
meshed reticulum formed of spines connected by
ridges (Figs 3C and 4D).

Habitat: On dead wood of conifers, in Abies-
Pinus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, &
August, 1989, 3250 m, ARP # 1147; 19 May, 1990, 3500 m, RP
# 1757.

Observations: Stemonitis nigrescens is an
unusual species which can be distinguished from
other species of Stemonitis by its almost black
sporangia and its spinose banded-reticulate
spores. There are other species of Stemonitis with
reticulate spores but only S. foliicola Ing has
rows of spines connected by ridges. That species
is distinguished from Stemonitis nigrescens by its
smaller sporangia and spores (8-9 um) and its
particular habitat on dead leaves (Nannenga-
Bremekamp, 1991).

Stemonitis pallida Wingate

Figs. 4E-G.

Habitat: On decayed wood of conifers and
living mosses, Abies-Pinus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 16 Sept.,
1988, 3300 m, RP # 490; 22 August, 1990, RP # 1830.

Observations: The distinctive features of
this species are the shortly stalked and crowded
sporangia (Fig. 4E), the peripheral spinose and
fragile net which dissipates at the apex and the
warted spores (Fig. 4F-G). Both of the specimens
from Tlaxcala agree with Nannenga-Bremekamp’s
(1991) description, although they have slight
differences in the ornamentation of the spores,
being taller in one of them (RP-490). However,
this variation falls within the range of the
species (Nannenga-Bremekamp, 1991). This species
usually grows on decaying wood but one of our
specimens (RP-1830) was found on living mosses.
Stemonitis pallida is close to S. herbatica which
is frequently found on living plants, but this

wat
Fe &

A. Schenella

germinal pore). B. Stemonitis hyperopta. C.

Stemonitis nigrescens.

Fig. 3. Spores of some Mexican Stemonitales.

simplex, (p

93

species has a complete surface net without spines.
Stemonitis flavogenita is also similar but it
usually has an apical membranous expansion in the
columella.

Stemonitis splendens Rost.

Habitat: On decayed wood and bark of
conifers, mosses, lichens and fungus, in Abies-
Pinus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 8 July,
1988, 3200 m, GF # 121; 13 July, 1988, 3100 m, HC # 138;
HC # 143; 14 July, 1988, 3400 m, RP # 182; 3500 m, HC #
160, RP # 158; 31 July, 1988, 3350 m, GF # 246; GF # 249;
RP # 245; 3450 m, RP # 236; 11 August, 1988, 3250 m, GF #
347; 12 August, 1988, 3400 m, RP # 334; 3500 m, RP # 304;
1 Sept., 1988, 3150 m, RP # 374; 2 Sept., 1988, 3300 m, RP
# 403; RP # 407; 3350 m, RP # 391; 15 Sept., 1988, 3300 m,
RP # 498; 3500 m, RP # 423; RP # 430; 16 Sept., 1988, 3150
m, RP # 526; 3100 m, HC # 628; 3200 m, RP # 508; 27 sept.,
1988, 3300 m, RP # 616; RP # 617; 3450 m, RP # 596; 3500
m, RP # 549; RP # 552; 04 Nov., 1988, 3300 m, RP # 684; RP
# 689; 26 Nov., 1988, 3250 m, GF # 795; GF # 803; 3300 m,
RP # 772; 3350 m, GF # 827; 19 Dec., 1988, 3300 m, RP #
840; 20 Jan., 1989, 3100 m, RP # 927; 10 March, 1989, 3400
m, RP # 971; 07 Jul., 1989, 3450 m, RP # 1071; 21 Jul.,
1989, 3350 m, GF # 1711; 18 August., 1989, 3250 m, GF #
1760; 3350 m, GF # 1763, GF # 1764; 31 August., 1989, 3300
m, RP # 1189; 3350 m, GF # 1118; 20 Oct., 1989, 3250 m,
GF # 458; 11 Jan., 1990, 3250 m, GF # 1230; 3300 m, RP #
1349; RP # 1357; 3500 m, RP # 1414; 09 Feb., 1990, 3300 m,
RP # 1460; 3350 m, HC # 1734; 3500 m, RP # 1497; 09 March,
1990, RP # 1556; 19 May, 1990, 3450 m, RP # 1742; 3500 m,
RP # 1759; 13 Jun., 1990, 3350 m, GF # 1564; GF # 1565.
el Escondido, Tlaxco Municipality, 30 August., 1989, GF

1080.

Observations: Stemonitis splendens is a
highly variable species, ranging from 5 mm to 25
mm in length, with a polygonal and fine or very
open mesh, from (10-)20 wm to 100(-125) pum,; with
dark colors, from black to dark purplish brown, or
sometimes reddish brown when the spores have
dispersed (Martin & Alexopoulos, 1969; Farr, 1976;
Lakhanpal & Mukerji, 1981). In its typical
expression it forms crowded sporangia growing on

94

a silvery hypothallus, making its determination
easy.

The Mexican specimens do not have the typical
silvery hypothallus of the species and many of
them are small and sometimes have expansions in
the capillitium and the apex of the columella.
These features point to S. flavogenita Jahn,
however, S. splendens can also have membranous
expansions in the capillitium (Martin &
Alexopoulos, 1969). When this happens, the
separation of the two species can be based on the
smaller size of the mesh and the paler color and
more delicate ornamentation of the spores of S.
flavogenita (Martin & Alexopoulos, 1969).
Nevertheless, in our specimens, all of the
combinations between sporangium size, mesh size,
columella expansion and kind of ornamentation can
be found, i.e. large sporangia with a small and
delicate mesh, expanded columella and either
strong or delicate ornamentation of the spores, or
small sporangia with open mesh, sometimes with
expansions in the columella and spores either
strongly or faintly marked.

Lakhanpal & Mukerji (1981) considered S.
splendens a variable species in its habit, height
of the sporangia and epispore, and they have
delimited it because of its metallic iridescence,
the attenuate coiled columella, and its small
superficial mesh. In some of the specimens from
Tlaxcala, we find iridescent sporangia, but this
feature is not constant. On the other hand,
Stojanowska (1983) and Bersan (1990) considered
that S. splendens develops on different kinds of
substratum, while S. flavogenita grows
predominantly’ on litter, and they pointed out that
the morphological characteristics are more
constant in the latter species.

We think that the specimens from. the
Malintzin Volcano belong to two or more species
which are difficult to separate using only
morphological features. We prefer to consider all
of them S. splendens until techniques are
developed to distinguish the species of this
complex.

95

Lamproderma columbinum (Pers.) Rost.

Habitat: On leaves of liverworts and mosses
Symblepharis vaginata (Hook) Wijk & Morgan,
Campylopus fragilis (Brid.) BSG and Brachithecium
sp., in Abies-Pinus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzi Volcano, 27 May,
1988, GF # 28; GF # 37; GF # 39; 23 June, 1988, RP # 41;
27 June, 1988, RP # 61; 12 August, 1988, 3450 m, GF # 379;
16 Sept., 1988, 3250 m, GF # 579; HC # 567; HC # 578; HC #
583; 27 Sept., 1988, 3250 m, HC # 755; HC # 757; 28 Sept.,
1988, 3250 m, HC 743; HC # 747; 4 Nov., 1988, 3250 m, GF
# 731; GF # 736; 3450 m, HC # 819; 26 Nov., 1988, 3250 m,
GF # 805; GF # 806; 19 Dec., 1988, 3250 m, GF # 866; GF #
869; 20 Jan., 1989, 3250 m, GF # 911; 10 Feb., 1989, 3250
m, GF # 939; 10 March, 1989, 3400 m, HC # 1340; 21 April,
1989, 3250 m, GF # 1018; GF # 1020; HC # 1367; 20 Oct.,
1989, 3250 m, HC. # 1519; HC # 1554; 11 Jan., 1990, 3250 m,
GF # 993; GF # 1241; GF # 1248; HC # 1602; HC # 1604; 3300
m, GF # 1256; 9 Feb., 1990, 3250 m, GF # 1334; GF # 1335;
9 March, 1990, 3250 m, HC # 1784; HC # 1789; HC # 1800; HC
# 1822; HC # 1823; HC # 1826; 23 March, 1990, 3250 m, GF #
1428; GF # 1435; GF # 1439; 19 April, 1990, 3250 m, GF #
1488; GF # 1498; 13 June, 1990, 3450 m, RP # 1789.

Observations: The distinctive features of
this species are the long stalks up to 4 mm, and
the brown or smokey color, the spinose or
verruculose ornamentation and the size (from 10 to
14 pm) of the spores. The Mexican specimens match
the description of Martin & Alexopoulos (1969),
except in the larger spores (10-18 4m).
Lamproderma columbinum is very close to lL.
echinulatum (Berk. ) Rost., but can be
distinguished by its larger spores and the longer
spines (up to 1.5 um). Lamproderma nigrisplendidum
Poelt was described from Nepal (Kowalski, 1970),
and is similar to L. columbinum except in the size
of the spores (14-17 pum). Nevertheless, UL.
nigrisplendidum is known only from the type
collection and it has been considered to be a
large-spored form of L. columbinum (Kowalski,
1970). If it is a good species, then the specimens
from Tlaxcala agree with L. nigrisplendidum
because of the size of their spores.

Lamproderma scintillans (Berk & Br.) Morgan

Habitat: On dead leaves of angiosperms in a
riparian mixed hardwood forest.

Specimens studied: Yauquemehcan Municipality,
Atlihuetzia Falls, 4 August, 1988, HC # 283; 5 August,
1988, HC # 288; RP # 251; RP # 264; 20 July, 1989, RP #
1076. :

Comatricha laxa Rost.

Habitat: On decayed wood of conifers and
mosses, in Abies-Pinus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzi Volcano, 16 Sept.,
1988, 3200 m, HC # 604; HC # 596; 28 Sept., 1988, 3200 m,
HC # 764; HC # 770; 04 Nov., 1988, 3500 m, HC # 805; 19
Dec., 1988, 3500 m, HC # 999; 19 May, 1989, 3200 m, HC #
1384; 27 Sept, 1989, 3500 m, HC # 686; HC # 687; 20 Oct.,
1989, 3250 m, GF # 1161; 3500 m, HC # 1582; 09 March,
1989, 3150 m, RP # 1559; 23 March, 1990, 3400 m, HC #
1361; HC # 1923; 3500 m, HC # 478; 19 April, 1990, 3250 m,
GF # 1497.

Observations: In its typical expression, this
species has very distinctive reddish ovate
sporangia. However, C. laxa is a highly variable
species in size, form of the sporangia and length
of the stalks. Hadrkénen & Uotila (1983) indicated
that the presence of horizontal primary branches
is a very useful feature for distinguishing this
species. The specimens from the Malintzin Volcano
are also highly variable but most of our specimens
have conspicuous horizontal primary branches.

Comatricha aff. lurida Lister

Sporangia gregarious, stalked, globose, up to
1 mm in total height and 0.3 mm in diameter;
peridium evanescent; stalk black, cylindrical but
wider at the base, shining; hypothallus reduced;
columella black, short, reaching half way into the
sporangium and divided apically into’ stout
branches from which the capillitium arises, with
membranous expansions between the branches of the
columella; capillitium semi-open, anastomosed,
dark brown, with blunt free ends; spores dark
brown in mass, violaceous brown by transmitted
light, globose, verrucose, 10 ~ 11 um in diameter.

97

Habitat: On decayed coniferous wood, in
Abies-Pinus forest.

Specimen studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 30 July,
1988, 3100 m, HC # 216.

Observations: The specimen studied is very
close to C. lurida, however, it has membranous
expansions between the branches of the columella,
the free ends of its capillitium are concolorous,
and the spores are slightly larger.

C. elegans (Racib.) Lister has a similar
branching columella but differs in having a short
columella divided near the base of the sporangiun.

Martin & Alexopoulos (1969) and Farr (1976)
considered that there may be variants connecting
Cc. lurida and C. elegans and their differences may
only be the expression of two extreme forms. The
Mexican specimen may be one of these variants, but
the presence of membranous expansions in the
columella separates it from those two species.
Similar expansions are found also in C. aggregata
Farr, but this species forms very crowded groups
of sporangia forming pseudoethalia, and it has a
persistent peridiun.

Comatricha nigra (Pers.) Schroet.

Habitat: Mainly on dead wood, but also on
bark, in Abies-Pinus forest.

Specimens studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 3 July,
1988, 3500 m, HC # 268; 8 July, 1988, 3250 m, HC # 122; 31
July, 1988, 3350 m, HC # 231; 3500 m, HC # 233; 31 August,
1988, 3350 m, HC # 1451; 1 Sept., 1988, 3150 m, RP # 370;
3500 m, HC # 486; 15 Sept., 1988, 3500 m, HC # 1873; 16
Sept., 1988, 3100 m, GF # 593; 4 Nov., 1988, 3350 m, GF #
707; 3500 m, RP # 736; 10 March, 1989, 3350 m, RP # 973;
21 April, 1989, 3500 m, HC # 1925; 20 Oct., 1989, 3350 m,
HC # 1506; GF # 1169; 3400 m, HC # 1583; 11 Jan., 1990,
3300 m, RP # 1331; RP # 1382; 9 Feb., 1990, 3100 m, RP #
1423; 3300 m, RP # 1450; 3500 m, GF # 1403; 9 March, 1990,
RP # 1545; RP # 1555; 19 April, 1990, 3450 m, ARP # 1713.

98

Comatricha rubens A. List.

=Collaria rubens (A. List.) Nann.-Brem.

Sporangia scattered to gregarious, up to 1 mm
in total height, globose to obovate, 0.3-0.5 mm in
diameter, pinkish brown; hypothallus membranous,
fragile, dark reddish, forming a disk at the base
of each individual sporangium; stalk black,
shining, cylindrical, approximately half the total
height; evanescent peridium except at the base of
the sporangium where it persists as a collar;
columella reaching to the center of the
sporangium, rigid, blunt; capillitium open,
arising along the length of the columella, bright
brown in transmitted light, formed by almost
parallel threads, with free, acute and paler ends;
spores pinkish brown in the mass, pale brown by
transmitted light, globose, spinulose, 7 - 8 pm in
diameter.

Habitat: On dead stems of herbaceous plants,
in Abies-Pinus forest.

Specimen studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano. 11 Jan.,
1990, 3350 m, RP # 1384.

Observations: The pinkish color and the
remains of the capillitium forming a collar are
the diagnostic features of this species. Farr
(1976) stated that it is unknown from the tropics;
however, she found a comparable specimen from
Mendoza, Argentina, which has smooth and slightly
larger spores. Nannenga-Bremekamp (1991) treated
this species in the genus Collaria.

Comatricha subcaespitosa Peck

=Stemonitopsis subcaespitosa (Peck) Nann.-
Brem.

Figs. 4H-I.

Habitat: On burned wood of conifers, in
Abies-Pinus forest.

Specimen studied: STATE OF TLAXCALA: Ixtenco
Municipality, east slope of the Malintzin Volcano, 28 Sept.,
1988, 3150 m, RP # 645; 26 Nov, 1988, 3250 m, RP # 758; 20
Jan, 1989, 3500 m, RP # 882; 9 Feb., 1990, 3250 m, RP #
1439; 3350 m, RP # 1467; 3400 m, RP # 1478; 3500 m, GF #
1412; 23 March, 1990, 3450 m, RP # 1630; 3500, RP # 1648;
13 Jun., 1990, 3400 m, RP # 1780.

99

Fig. 4. A-B. Stemonitis hyperopta (RP-127). A. Spores in
surface view (Nomarski Interference Contrast: NIC X 1100);
B. Spores in optical section (NIC X 1100). C-D Stemonitis
nigrescens (RP- 1757); C. Sporangia (X 8.8); D. Spores (NIC
X 1100). E-G. Stemonitis pallida (RP-490); E. Sporangia (X
13.2); F. Spores in surface view (NIC X 1100); G. Spores in
optical section (NIC X 1100). H-I. Comatricha subcaespitosa

(RP-758); H. Superficial net of the sporangia (xX 110); I.
Spores (NIC X 1100).

100

Observations: Our specimens agree with
descriptions of Martin & Alexopoulos (1969) and
Nannenga-Bremekamp (1991) (as Stemonitopsis
subcaespitosa) except in the color of the
hypothallus, being reddish brown and not silvery
as those workers indicated. Nevertheless, the
evident membranous expansions in the internal net,
the almost complete with angular meshes of the
surface net (Fig. 4H) and the warted spores (Fig.
41) distinguish this species from others in the
genus. Superficially, Comatricha subcaespitosa
looks like a species of Stemonitis because of its
almost complete peripheral net, but it can be
distinguished from members of that genus by the
structure of its stalk which is solid and fibrous.

Comatricha tenerrima (Curt.) List.

Habitat:On dead wood of angiosperms and stems
of herbaceous plants, in Pinus~ Quercus forest and
riparian mixed hardwood forest.

Specimens studied: STATE OF TLAXCALA: Totolac
Municipality, Tepeticpac Hill, 26 July, 1991, HC # 1996.
Yauquemehcan Municipality, Atlihuetzia Falls, 4 August,
1988, HC # 282.

DISCUSSION

Both families of Stemonitales recognized by
Martin et al. (1983) and Nannenga-Bremekamp
(1991), were represented in the State of Tlaxcala:
Schenellaceae and Stemonitaceae. Species found in
this geographical area belong to nine of the 16
genera of Stemonitales (Nannenga-Bremekamp, 1991).
Stemonitis and Comatricha are the most frequently
collected genera and have the highest number of
species. On the other hand, Schenella, Colloderma,
Diacheopsis, Collaria and Enerthenema were
represented each by only one species. Also, it is
important to emphasize the infrequency of the
genus Lamproderma, with only two species in the
state, and only one in the main area of study. All
of the recorded species in this paper were
previously unknown to Tlaxcala State, three
species for the Neotropics and two species for
Mexico,

101

The species of Stemonitales from Tlaxcala
represent around 12.3 % of the total number of
known species of this Order (Nannenga~Bremekanp,
1991), and around 58.3 % of the known Stemonitales
from Mexico (Trujillo, 1988; Ogata et al., 1994).
We add two genera of Myxomycetes previously
unrecorded from Mexico: Schenella and Diacheopsis
(Braun & Keller, 1986; Ogata et al., 1994). Now 45
of the 59 known genera of Myxomycetes are recorded
from Mexico. Braun and Keller (1986) listed 164
species of Myxomycetes collected from Mexico up to
1986. Since that date, 39 taxa have been added to
that list (Trujillo et al., 1986; Trujillo, 1988;
Villarreal, 1990; Hernandez Cuevas et al., 1991;
Ogata et al., 1994), including the taxa recorded
in this paper. The number of recorded species from
Mexico is now 203 which is about 33.8 % of the 600
described taxa of Myxomycetes (Nannenga-Bremekamp,
1991).

ACKNOWLEDGEMENTS

We wish to thank Mr. D. W. Mitchell, Dr. H. W Keller,
Mr. K. L. Braun and Dr. C. Ladd for revising this
manuscript. Mr. D.W. Mitchell and Mrs. Nannenga-Bremekamp
helped identify some of our specimens.

LITERATURE CITED

Acosta, R. & A. Kong, 1991. Guia de fas excursiones
botdnicas y micoldégicas al cerro de E] Pefion y Cafiada
Grande del estado de Tlaxcala. IV Congreso Nacional de
Micologia. Universidad Auténoma de Tlaxcala, Sociedad
Mexicana de Micologia y Jardin Botanico de Tizatlan.
Tlaxcala. 82 pp.

Bersan, F., 1990. I1 genere Stemonitis Roth (Myxomycetes;
Stemonitomicetidae) in Italia. AWB. 33:210-216.
Braun, K. L. & H. W. Keller, 1976. Myxomycetes of Mexico I.

Mycotaxon 3:297-317.

Braun, K. L. & H. W. Keller, 1986. Myxomycetes of Mexico
III. Rev. Mex. Mic. 2:25-39

CETENAL, 1975 A. Carta edafoldégica, Tlaxcala, 1:250000.
S.P.P., México, D.F.

CETENAL, 1975 B. Carta climdtica, Tlaxcala, 1:250000.
S.P.P., México, D.F.

102

Farr, M. L., 1976. Myxomycetes. in: Flora Neotropica. The
New York Botanical Garden. Nueva York. 304 pp.
Galindo, F. G., 1992. Algunos hongos del Jardin Botanico

Tizatldn, Gobierno del estado de Tlaxcala.

Harkoénen, M. & P. Uotila, 1983. Turkish myxomycetes
developed in moist chamber cultures. Karst. 23:1-9.

Hernandez-Cuevas, L., M. Rodriguez-Palma, G. Galindo-F lores
& A. Estrada-Torres, 1991. New records of myxomycetes
from Mexico. Mycotaxon 62:17-27.

Keller, H. W. & K. L. Braun, 1977. Myxomycetes of Mexico II.
Bol. Soc. Mex. Mic. 11:163-180.

Kowalski, D. T., 1970. The species of Lamproderma.
Mycologia 62:621-672.

Kowalski, D. T., 1975. The genus Diacheopsis. Mycologia
67:616-628. |
Lakhanpal, T. N. & K. G. Mukerji, 1981. Taxonomy of Indian

Myxomycetes. Cramer. Voduz, 531 pp.

Lé6pez A., L. Villarreal & A. Sosa., 1981. Estudio sobre los
myxomycetes del estado de Veracruz, III. Bol. Soc.
Mex. Mic. 16:77-93.

Martin, G. W. & C. J. Alexopoulos, 1969. The Myxomycetes.
Univ. Iowa. Academic. Press. Iowa City, 561 pp.
Martin, G. W., C. J. Alexopoulos & M. L. Farr. 1983. The
Genera of Myxomycetes. Univ. Iowa. Academic. Press.

Iowa City, 102 pp.

Mitchell, D. W., 1995. Synkey [Synoptic key to all described
Myxomycetes for IBM Personal Computer].

Moreno, G., C. Illiana & M. Heykoop., 1989. Contribution to
the study of the Myxomycetes in Spain I.
Mycotaxon 34; 623-635.

Munsell, 1992. Munsell soil colour charts. Kollmorgen
Corporation, Baltimore.

Nannenga-Bremekamp, N. E., 1991. A guide to temperate
Myxomycetes. Biopress. Bristol. 409 pp.

Ogata, A., D. Nestel, V. Rico-Gray & G. Guzman., 1994. Los
Myxomycetes citados de Mexico. Acta Bot. Mex. 27:39-
51

Stojanowska, W., 1983. Myxomycetes of the litter. Acta
Mycol. 19:21-30.

Trujillo, F. I., 1988. Contribucién al conocimiento de los
myxomycetes de la Sierra de Manatlan, Jalisco. Tiempos
de Ciencia. 12:20-27.

Trujillo, F. I., M. M. Castafieda & L. Guzman-Davalos, 1986.
Hongos del estado de Jalisco, IV. Los myxomycetes
conocidos. Tiempos de Ciencia 5:42-51.

Villareal, L. 1990. Estudio sobre los myxomycetes de Mexico
I. Nuevos registros. Mycol. Neotrop. Ap]. 3:67-79.

MYCOTAXON

Volume LX, pp. 103-110 October-December 1996

STUDIES ON MEDITERRANEAN HYPHOMYCETES. III.
QUADRACAEA MEDITERRANEA
ANAM.-GEN. AND SP. NOV.

LUNGHINI D., PINZARI F. AND ZUCCONI L.*

Laboratorio di Micologia, Dipartimento di Biologia Vegetale,
Largo Cristina di Svezia 24, I-00165 Roma, Italy
* Dipartimento di Scienze Ambientali, Via San Camillo de Lellis,
I-01100 Viterbo, Italy

Abstract

Quadracaea mediterranea anam.-gen. et sp. nov., an
hyphomycete recorded on leaves of Quercus ilex L. from central
Italy, is described, discussed and illustrated, also by means of
SEM.

Keywords: Hyphomycetes, Quadracaea_ mediterranea,
Endophragmiella, Selenosporella, Italy, new genus

During mycological studies carried out in different sites of central
Italy, characterized by Mediterranean shrubby vegetation (Lunghini, 1994,
1995; Zucconi et al., 1995), an interesting fungus (dematiaceous
Hyphomycete) was collected on Quercus ilex L. leaf litter.

Quadracaea anam.-gen. nov.

[Etym.: dedicated to the memory of the mycologist Livio Quadraccia
(1958-1993), University "La Sapienza" of Rome]

Coloniae effusae. Mycelium partim superficiale, partim immersum,
ex hyphis septatis ramosisque compositum. Hyphopodiis nullis.
Conidiophora macronemata, mononemata, erecta, septata, crassitunicata.
Cellulae_conidiogenae integratae, terminales, postea intercalares,
polyblasticae, cellulas separantes acropleurogenas producentes.
Subsequentes cellulae conidiogenae proliferatione percurrente oriuntur ex

104

septo basilari cellulae terminalis. Cellulae separantes ampulliformes, ad
apicem post rhexolyticam secessionem conidiorum apertae, vacuae.
Conidia blastica, solitaria, cicatrice basilari margine fimbriata praedita.
Species typica: Quadracaea mediterranea anam.-sp. nov.

Colonies on natural substratum effuse. Mycelium superficial or
immersed, composed of branched, septate hyphae. Hyphopodia absent.
Conidiophores macronematous, mononematous, straight, septate, thick-
walled. Conidiogenous cells integrated, terminal, becoming intercalary,
polyblastically producing acropleurogenous groups of separating cells.
Successive percurrent proliferations occur at the basal septum of the
terminal separating cell into and beyond it. Separating cells ampulliform,
with an open end appearing empty after rhexolytic conidial secession.
Conidia blastic, solitary, bearing a frill at the base.

Quadracaea mediterranea anam.-sp. nov. (Fig. 1)

Coloniae amphigenae, effusae, brunneae usque ad atro-brunneae,
pilosae. Mycelium partim superficiale, partim in substrato immersum, a
luteo ad brunneum, ex hyphis septatis, ramosis, levibus, usque ad 4 um
latis, ad conidiophororum basim inflatis compositum. Conidiophora
solitaria vel (3-4) fasciculata, recta vel flexuosa, levia, luteo-brunnea,
sursum pallidiora, ad basim lobatam 6-11.5 wm lata, usque ad 250 um
longa et in parte inferiore 4-7 [um lata, cellulas separantes in partibus
variis ferentia. Cellulae_ conidiogenae integratae, terminales, deinde
intercalares post percurrentem proliferationem, subcylindricae, 5.7-9 x
3.6-5 tum, colore conidiophoris similes, polyblasticae, cellulas separantes
proferentes. Cellulae separantes acropleurogenae, simplices vel usque ad 7
verticillatae, ampulliformes, apicem versus attenuatae, post rhexolyticam
secessionem conidii apertae, vacuae, tenuitunicatae, leves, pallide
brunneae, 4.5-6.7 x 3.5-4.4 um. Raro ramus e latere per proliferationem
percurrentem emergens. Conidia Solitaria, sicca, subovata ad
obpiriformia, 3-(raro 4-) septis transversalibus praedita, modice ad septa
constricta, 20-28.5 x 8.2-12 um; cellula basilaris fere crassitunicata, levis,
dilute brunnea, cicatrice basilari margine fimbriata praedita; secunda et
tertia cellulae valde crassitunicatae, leves (paulo asperata si SEM
observantur ), atro-brunneae, secunda cellula reliquis amplior et obscurior;
cellula apicalis tenuitunicata, levis, subhyalina, emergentia subconica
apicali praedita, quae phialoconidia gignere potest; phialoconidia
fusiformia, leviter falcata, aseptata, hyalina, Selenosporellae similia, 5.5-
8.8 x 0.8-1.2 pum. .

In foliis emortuis Quercus ilicis L., in loco "Riomarina", in Insula
Elbae (Prov. Livorno, in Italia), legit D. Lunghini, 8.IV.1989, IMI
367816, holotypus.

105

Fig. 1. Quadracaea_ mediterranea: a and b, mature and young
conidiophores; c, detail of conidia; d, colony on its natural substratum.

106

Colonies amphigenous, effuse, dark brown to blackish brown, hairy.
Mycelium partly superficial, partly immersed in the substratum, composed
of branched, smooth, septate, light brown or yellowish brown, up to 4 um
wide hyphae, swollen near the conidiophore origin. Conidiophores arising
singly or in small groups (3-4), straight or flexuous, smooth, dark
yellowish brown, paler towards the apex, with a lobate base measuring 6-
11.5 um in width, up to 250 um long and 4-7 um wide near the base,
bearing separating cells at different levels. Conidiogenous cells
subcylindrical, 5.7-9 x 3.6-5 um, concolorous with the conidiophore,
polyblastic, producing apical and lateral separating cells, integrated,
terminal, becoming intercalary by percurrent proliferations which arise
by an extension of the basal septum of the apical separating cell into and
beyond it. Separating cells acropleurogenous, simple or in clusters when
numerous (up to 7), ampulliform, tapered at the apex, empty, leaving an
open end after the rhexolytic conidial secession, thin-walled, smooth, pale
brown, 4.5-6.7 x 3.5-4.4 um. Rarely, an occasional lateral percurrent
proliferation may produce a lateral branch. Conidia solitary, dry, suboval
to obpyriform, 3-(rarely 4-) transversely septate, slightly constricted at
the septa, 20-28.5 x 8-12 um; basal cell slightly thick-walled, smooth, pale
brown, bearing a conspicuous frill at the base; second and third cells very
thick-walled, smooth at the light microscope (finely roughened under
SEM), very dark brown, the second broader and darker than the others;
apical cell thin-walled, smooth, subhyaline, bearing a subconical
elongation which can produce phialoconidia. Synanamorphic phialoconidia
fusiform, slightly falcate, aseptate, hyaline, produced in a Selenosporella-
like pattern, 5.5-8.8 x 0.8-1.2 um. An attempt to obtain this species in
pure culture was unsuccessful. No ascocarps of a possible teleomorph were
found.

Other material examined: on a leaf litter of Quercus ilex L., at "San
Liberato" (Terni, Italy), legit D. Lunghini, 29.1V.1993, IMI 367817,
paratypus; on leaf litter of Q. ilex, at "Macchiagrande" (Focene, Rome),
legit F. Pinzari, 23.X1.1992, ROHB 181, paratypus; on leaf litter of Q.
ilex, at "Cala Martina" near Follonica (Grosseto, Italy), legit L. Zucconi,
18.X.1993, ROHB 179, paratypus; on leaf litter of Q. ilex, at "Alma"
(Grosseto, Italy), legit L. Zucconi, 18.X.1993, ROHB 180, paratypus.

Quadracaea can be distinguished from other similar hyphomycete
genera by the combination of polyblastic conidiogenous cells, each
producing more ampulliform separating cells, rhexolytic conidial
secession and apical percurrent proliferation.

Conidial production and percurrent proliferation (Fig. 2) are similar
with those of Endophragmiella boewei (Crane) S. Hughes (1979). In
Quadracaea the septa at the base of the separating cells are formed after
those at the base of the conidia at a time when the maturing conidia
become pigmented. This process leads to the appearance of verticillate
pigmented empty cells, which we prefer to call separating cells while

107

Fig. 2. Quadracaea mediterranea. Diagrammatic representation of
delimitation of separating cell, conidium secession and origin of
percurrent proliferation from terminal conidiogenous cell.

reserving the term conidiogenous cells to the cells of one level below.
They are comparable to the "terminal cells" described by Hughes (1979)
for E. boewei. The conidiogenous cells are therefore integrated in the
main axis of the conidiophore. They produce separating cells
polyblastically, the first one apically and the successive ones laterally.
Lateral separating cells, generally in clusters, are produced near the basal
septum of the apical separating cell (acropleurogenously) and only
occasionally at lower levels. Rarely, branches are produced by percurrent
proliferations through lateral separating cells.

The verticillate disposition of the separating cells observed in
Quadracaea superficially resembles that of branches and conidiogenous
cells in Physalidiopsis Castafieda & Kendrick (1990). However, according
to Castafieda (pers. comm.), the conidiophores in Physalidiopsis are
definitely branched and the conidia are stauroconidia. Lateral branches are
also present in the apparently similar Endophragmiella verticillata S.
Hughes (1978).

Quadracaea has, in common with the above reported genera, the
occurrence of synanamorphic Selenosporella-like phialoconidia.

Figs 3 and 4 show photographs of Q. mediterranea by light and
scanning electron microscopy. The polyblastic conidiogenous cells produce

Fig. 3. Quadracaea mediterranea: a and c, conidia and conidiophore apex
(a x 850; c x 1600); b, lateral production of branches (x 950); d, conidium

with basal frill and apical synanamorphic phialoconidia (x 1900).

109

Fig. 4. Quadracaea mediterranea: a, conidiophores on the leaf surface (bar
= 10 um); b and d, polyblastic conidiogenous cells and empty separating
cells (bar = 1 um); c and g, conidia and separating cells (c, bar = 5 um; g,
bar = 1 um); e and f, synanamorphic phialoconidial production at the
conidial apex (bar = 1 um).

110

several ampulliform separating cells (Fig. 4 b, d), which leave an open end
after rhexolytic conidial secession and appear empty (Fig. 4 d, g). Up to
seven separating cells disposed in a cluster have been observed, but the
complexity of the fertile region could probably be influenced by
environmental and nutritional conditions. Percurrent proliferation occurs
through the apical separating cell (Fig. 3 a-c) and this mode of
conidiogenesis can be repeated several times. Rarely proliferation near the
basal septum (Fig. 3 b) or occasionally inside lateral separating cells (Fig.
1 a) may produce a branch. Conidia generally remain attached to the
separating cells and the result of successive proliferations is a complex
structure (Fig. 4 c). Conidia are smooth-walled when observed under the
light microscope (Fig. 3 a-d), but the two central cells are found to be
lightly roughened under SEM (Fig. 4 c); they are provided with a frill at
the closed base due to rhexolytic liberation (Fig. 4 g).

Detached conidia still connected with the separating cells at the base
have occasionally been observed.

The authors wish to thank Prof. Walter Gams, Dr Rafael Castafieda
Ruiz and Prof. Silvano Onofri for kindly reviewing the manuscript and
the Museo Provinciale di Storia Naturale (Livorno) for the invaluable
cooperation offered. This research was supported by a grant from
M.U.R.S.T. (Ministero dell'Universita e della Ricerca Scientifica e
Tecnologica).

References

Castaneda Ruiz, R.F. & Kendrick, B. (1990). Conidial fungi from
Cuba: I. Univ. Waterloo Biol. Ser. 32: 53 pp.

Hughes, S.J. (1978). Endophragmiella verticillata. Fungi Canadenses
No. 130.

Hughes, S.J. (1979). Relocation of species of Endophragmia auct. with
notes on relevant generic names. N. Z. J. Bot. 17: 139-188.

Lunghini, D. (1994). Menispora convoluta, a new dematiaceous
Hyphomycete from central Italy. Mycotaxon 50: 429-433.

Lunghini, D. (1995). Studi sugli Ifomiceti mediterranei. II. Ifomiceti
Demaziacei dell'Isola d'Elba (Livorno). Nota preliminare. Quaderni
Mus. St. Nat. Livorno (in press).

Zucconi, L., Facchin, M. & Perini, C. (1995). Microfungal
communities in Mediterranean evergreen forests (Central Italy).
Bocconea (in press).

MYCOTAXON

eS y-+-eae
Volume LX, pp. 111-123 October-December 1996

HETEROBASIDIOMYCETES FROM MAJORCA &
CABRERA
(Balearic Islands)

PETER ROBERTS

The Herbarium, Royal Botanic Gardens, Kew, Surrey, TW9 3AE, England
E-mail: P.Roberts@rbgkew.org.uk

Seven species of the Platygloeales, Tremellales, and Tulasnellales are described and
illustrated. The mycoparasitic genus Sp/iculog/oea, provisionally assigned to the
Platygloeales, is described as new, based on the new species Sp/culog/oea occulta.
Tulasnella balearica is described as new. Basidiodendron nodosum, B. rimulentum,
Ceratosebacina longispora, Sebacina epigaea, and Serendipita vermifera are
described and illustrated. Eleven additional heterobasidiomycete species, all from
Majorca, are documented.

A number of new and unusual heterobasidiomycetes were collected
during the British Mycological Society's 1992 Overseas Foray to the
Mediterranean islands of Majorca and Cabrera, part of the Balearic group
lying off the east coast of Spain.

The foray, held in November, was based at Port de Pollenca on the
north-east coast of Majorca, with most collecting undertaken in the
adjacent coastal strip and in the highlands of the Serra de Tramuntana (ca
500-1200 m). A single short visit was made to the small island of Cabrera
some 10 km south of Majorca. Despite previous rain, weather during the
foray was rather dry so that few of the more conspicous
heterobasidiomycetes were in evidence, most collections being from fallen
wood and other material in damp litter.

New and less common collections are described in detail below,
followed by a checklist of additional heterobasidiomycetes collected
during the foray.

bi2

PLATYGLOEALES

Spiculogloea P. Roberts gen. nov.

Basidiomata nulla, in hymeniis fungorum parasitica. Hyphae angustae, in typo
generis fibulatae, haustoria tremelloidea ferentes. Basidia cylindracea, transeptata,
e probasidiis exorientia; plus minusve crassitunicata, cyanophila, subtiliter spiculata
vel granulata. Conidia in typo generis oblonga vel subcylindrica, e conidiophoris
simplicibus exorientia. Sporae in typo generis fusiformes.

Typus generis: Spiculogloea occulta

Basidiome within the hymenium of the host; not visible to the naked eye.
Hyphae narrow, clamped in the type species, connected to the host
hyphae by tendril-like haustorial cells, similar to those found in the genus
Tremella Pers. Basidia tubular, laterally septate, arising from probasidia;
both probasidia and basidia somewhat thick-walled, cyanophilous, and
covered with fine spicules or granules. Conidia ellipsoid in the type
species, arising in clusters Gn simple conidiophores produced alongside
the basidia. Spores fusiform in the type species.

The new genus Spiculogloea is distinguished by its _ parasitic,
intrahymenial habit, the presence of 7reme//a-like haustorial cells, and the
curious, cyanophilous, finely spiculate or granular basidia arising apically
from small probasidia. A second, much smaller, narrow-spored species
has been found in England as a_e parasite of resupinate
heterobasidiomycetes.

Two other auricularioid genera are known to possess similar haustorial
cells: Occultifur Oberw. and Zygogloea P. Roberts. The former is a
parasite of Dacrymyces Nees: Fr., and has smooth, thin-walled basidia
which do not arise from probasidia. The latter is a parasite of Myxarium
nucleatum Wallr., has coiled basidia lacking probasidia, and produces
distinctive ‘twinned’ zygoconidia. Neither closely resembles the new
genus, Spiculogloea.

is

HOO 080004

Fig. 1. Spiculogloea occulta: basidiospores from print; basidia showing haustorial
cells, conidiophore and conidia, and swollen probasidia (K(M) 28706; holotype]

Spiculogloea occulta P. Roberts sp. nov.
(Fig. 1)

Basidiomata nulla, in hymeniis Corticiacearum parasitica. Hyphae angustae, 0.5-1.5
um latae, fibulatae, haustoria tremelloidea ferentes. Basidia_ cylindracea,
transeptata, stipitata, usque 40 um longa, e probasidiis parvis exorientia; plus
minusve crassitunicata, cyanophila, subtiliter spiculata vel granulata. Conjidia
oblonga vel subcylindrica, 3.5-5.0 x 2.0-2.5 wm, e conidiophoris simplicibus
exorientia. Sporae fusiformes, 3.5-7.5(-8.0) x 2.5-4.5(-5.0) um.

In hymenio Hyphodermatis argillacei, Son Moragues, Majorca, Balearicae Insulae,
11 Nov. 1992, P. Roberts 576, holotypus, K(M) 28706.

114

Basidiomes none, parasitic in the hymenium of resupinate, corticioid fungi.
Hyphae clamped, rather thin, 0O.5-1.5 um _ wide. MHaustorial cells,
subtended from clamp-connexions, arising singly or in clusters from the
hyphae. Bas/dia auricularioid, 4-celled, laterally septate, arising apically,
often on a distinct stalk of variable length, from small, clavate probasidia
which are clamped at the base; total length up to 45 um. Basidia and
probasidia somewhat thick-walled, cyanophilous, and covered in fine
spicules or granulations. Sterigmata sinuous, up to 15 um long, the
sterigma of the apical cell arising laterally. Conidia formed laterally, in
clusters, on simple tubular or weakly clavate conidiophores arising from
basidia-bearing hyphae; individual conidia apiculate, oblong to
subcylindrical, 3.5-5.0 x 2.0-2.5 um. Basidiospores widely fusiform, 3.5-
7.5(-8.5) x 2.5-4.0(-5.0) um from print, occasionally producing secondary
spores. Germination not seen.

Specimen examined: in hymenium of Hyphoderma argillaceum (Bres.)
Donk on fallen wood, Son Moragues, Majorca, 11 Nov. 1992, P. Roberts
576, holotype, K(M) 28706.

The common auricularioid parasite of corticioid fungi is Co/acogloea
peniophorae (Bourdot & Galzin) Oberw. & Bandoni, normally found on or
in Hyphoderma praetermissum (P. Karst.) J. Erikss. & A. Strid. C.
peniophorae can easily be distinguished from Spiculogloea occulta by its
smooth basidia, lack of haustorial cells, and ventrally flattened,
subellipsoid spores.

Whether Spiculogloea occulta is host specific is one of many
questions still to be answered. It is hoped that this initial description may
encourage corticiologists to specifically search for this and other
interesting hymenial parasites. They may be a more common and diverse
group than the comparatively few records suggest.

TREMELLALES

Basidiodendron nodosum Luck-Allen. Can. J. Bot. 41: 1045
(1963) (Fig. 2)

Basidiomes thin, effused, ceraceous, pale grey. Hyphae narrow,
agglutinated, and closely adhering to the substrate. Basidia 2-4 celled,
globose to ellipsoid, ca 8-15 um wide, tending to arise acropetally on the
sub-basidial hyphae, but not conspicuously so (compared to B. rimulentum
below). Sterigmata rather short, narrowing from base to tip. Cystidia

115

Fig. 2. Basidiodendron nodosum: a) basidiospores from print, basidia and cystidium
[K(M) 28708]; 6) [K(M) 28707]

tubular with lateral basal stalk; length variable, but normally equal to the
depth of the hymenium; not or barely projecting; some hyaline, most with
highly refractive, cyanophilous, yellowish-brown contents. Spores broadly
ellipsoid (Q = 1.1-1.4), 6.0-8.0 x 5.0-6.5 um (Roberts 623); 8.0-9.0 x
6.5-7.0 um (Roberts 583) from prints.

Specimens examined: on woody remains of Agave (?), Port de Pollenca,
Majorca, 12 Nov. 1992, P. Roberts 583, K(M) 28707; on fallen twig,
probably Erica multiflora, Cabrera, 15 Nov. 1992, P. Roberts 623, K(M)
28708.

Basidiodendron nodosum does not appear to have been reported from
Europe before, though it is not uncommon in southern England and is
probably widespread throughout the continent. Collections from the

116

former Soviet Union with similar, broadly ellipsoid spores were referred by
Wells & Raitviir (1975) to B. cinereum (Bres.) Luck-Allen, a closely
related taxon differing only (if at all) in its longer, ellipsoid or oblong
spores.

The Canadian type collection of B. nodosum (on decayed Pinus, Little
White River, Ontario, 14 Sep. 1956, R.F. Cain, TRTC 33408) has spores
of almost identical size and shape to the Cabrera collection cited above.
The Majorcan collection has slightly larger spores and rather larger, more
globose basidia, but is within the degree of variation seen in British
collections. The species, in common with the genus as a whole, appears
to be litter-inhabiting without any noticeable substrate preferences.

Fig. 3. Basidiodendron rimulentum: basidiospores from print, basidia and cystidium
[K(M) 28709]

Basidiodendron rimulentum (Bourdot & Galzin) Luck-Allen.
Can. J. Bot. 41:1037 (1963) (Fig. 3)

Basidiome thin, effused, ceraceous, pale grey. Hyphae narrow,
agglutinated, closely adhering to the substrate. Basidia 4-celled, rather
small, ca 3.5-4.0 um wide, arising acropetally on conspicuous, well-
defined, sub-basidial stalks. Sterigmata short. Cystidia tubular, similar to
those of B. nodosum above. Spores appearing ellipsoid in side view,
reniform or heart-shaped from above or below, 3.0-3.5(-4.0) x 3.5-5.0 um
from print.

Specimen examined: on fallen twig, probably Erica multiflora, Cabrera, 15
Nov. 1992, P. Roberts 624, K(M) 28709.

117

Basidiodendron rimulentum is a widespread but uncommon species, very
similar to B. eyrei (Wakef.) Luck-Allen which differs in having more
conventional, globose spores. Both species have well-developed and
conspicuous basidial ‘stalks' on which the basidia are produced in
succession. In larger-spored species, such as B. nodosum above, these
stalks are much less well-defined.

iD fe (bied a oS
10 um

Fig. 4. Ceratosebacina longispora: basidia and basidiospores [K(M) 26202]

Ceratosebacina longispora (Hauerslev) P. Roberts.
Mycological Research 97: 470 (1993) (Fig. 4)

Basidiome not visible to the naked eye in this specimen; when visible,
thin, effused, ceraceous, greyish. Hyphae clamped, ca 2.5-4.0 um wide,
not appearing gelatinized, forming a loose, irregular hymenium similar to
that seen in species of Ceratobasidium D.P. Rogers. Hyphidia absent.
Basidia 2-4 celled, subglobose to ellipsoid, ca 10 um wide, but often
appearing cuboid or snouted as a result of short hyphal outgrowths.
Sterigmata in this and the majority of specimens consistently short, wider
at the base than the apex. Spores elongated, narrowing at each end,
obtusely angled at mid-point, 23-33 x 2.5-3.0(-3.5) um from print.

118

Specimen examined: on fallen wood, Son Moragues, Majorca, 11 Nov.
1992, P. Roberts 574, K(M) 26202.

This inconspicuous but distinctive species has recently been redescribed
by Roberts (1993) from a number of British collections. The type
collection is from Denmark and specimens have been seen from Belgium,
France, and Norway, suggesting C. /ongispora is widespread in Europe.

Sebacina epigaea (Berk. & Broome) Neuhoff. Zeitschrift fur
Pilzkunde 15(3): 71 (1931) (Fig. 5)

Basidiomes at first pustular, subgelatinous, greyish blue; then coalescing
and appearing tuberculate, becoming ceraceous and opaque; finally
encrusting, uneven to smooth, subcartilaginous, greyish to ochraceous,
up to 10 cm or so across. Hyphae unclamped, ca 2.0-4.0 um wide, in a
gelatinous context, basal hyphae becoming somewhat thick-walled and
cyanophilous. Hyphidia weakly branched, forming a layer above the
basidia. Basidia 4-celled. subglobose to ellipsoid, often somewhat stalked
and occasionally myxarioid. Sterigmata sinuous, extending indefinitely to
the hymenial surface. Spores ellipsoid, 9.0-12.0 x 6.0-7.0 um (D.
Jefferies, Cabrera); 11.5-13.0 x 9.0-11.0 um (Roberts 559); 11.5-15.5 x
8.5-12.0 um (Roberts 558), from prints. Resting spores formed from
primary or secondary basidiospores, the spore content shrinking into a
thick-walled, refractive, cyanophilous central section, sometimes leaving
the outer remnants of the spore as hyaline, thin-walled projections.

Specimens examined: encrusting surface of half-buried wood, Menut,
Majorca, 10 Nov. 1992, P. Roberts 558, K(M) 26231; same date and
location, on damp compacted earth, P. Roberts 559, K(M) 26242;
encrusting fallen wood and litter, Cabrera, D. Jefferies, 15 Nov. 1992,
K(M) 26822.

Sebacina epigaea is a common, but seldom described species, with a
preference for growing on damp, compacted earth. In such fruit bodies,
the hymenium is non-geotropic, a feature shared with other species of
Sebacina Tul. sensu stricto.

The appearance of the fruit body changes markedly with age (see
description above). Furthermore the number and appearance of the resting
spores, a distinguishing feature of Sebacina epigaea, also seem to change
with age. In young fruit bodies (such as the Cabrera specimen) they are
normally sparse or absent, and appear to be derived entirely from primary
basidiospores. The spore contents shrink into a concentrated, refractive,
cyanophilous central section delimited by thickened walls; the outer part

119

of the spore simply collapses, so that the resting spore appears rather
angular. In older fruit bodies, the number of resting spores increases, and
most are derived from secondary spores. These are often highly irregular,
sometimes becoming stellate, with numerous hyaline projections. Often
the empty shell of the primary basidiospore (or at least the connecting
sterigma) remains attached. Very old fruit bodies have been found which
only contain resting spores, with no functional basidia remaining.

The two Majorca specimens cited above have a high proportion of
unusually large basidiospores, but otherwise appear typical.

Fig. 5. Sebacina epigaea: hyphidium and basidia; nodulose resting spores;
unclamped hyphae in context; basidiospores from print [K(M) 26242]

120

Serendipita vermifera (Oberw.) P. Roberts, Mycological
Research 97: 474 (1993) (Fig. 6)

Basidiome not macroscopically visible. Hyphae ca 1.0-2.0 um wide, not
forming an hymenium but trailing loosely over the substrate; septa not
seen. Basidia 2-4 celled, subglobose to ellipsoid, ca 6.0 um wide, arising
in clusters of 3-5 at intervals along the hyphae. Sterigmata very variable,
often branching off subsidiary sterigmata. Spores vermiform, straight or
slightly sinuous, 36-50 x 1.5-2.5 um from print.

Specimen examined: on fallen wood, with or near Botryobasidium
isabellinum (Fr.) D.P. Rogers, Bosc de Monnaber, Majorca, 13 Nov. 1992,
P. Roberts 587, K(M) 26201.

Oberwinkler (1963) originally described the species with scattered clamp
connexions, but Warcup & Talbot (1967), studying cultured isolates from
orchid roots, could find none. British collections appear unclamped
(Roberts, 1993).

10 pm

Fig. 6: Serendipita vermifera: clustered basidia; basidiospores from print
[K(M) 26201]

121

TULASNELLALES

Tulasnella balearica P. Roberts sp. nov. (Fig. 7)

Basidiomata effusa, ceracea, violaceogrisea. Hyphae non fibulatae, ca 3.0-4.0(-6.0)
um latae. Basidia clavata, ca 5.5-6.5 um lata. Sterigmata 4, ellipsoideae vel
fusiformes. Sporae angustae, 9.0-20.0(-26.5) x 2.5-3.0(-3.5) um (Q = 3.0-9.0).

In spica dejecta Zeae mayis, Binifaldo, Majorca, Balearicae Insulae, 10 Nov. 1992,
P. Roberts 565, holotypus, K(M) 28705

Fig. 7: Tulasnella balearica: basidia; basidiospores from print {K(M) 28705;
holotype]

122

Basidiomes effused, thin, ceraceous, violaceous-grey. Hyphae unclamped,
conventionally branched, ca 3.0-4.0 um wide, occasionally swollen up to
6.0 um wide in the hymenium. Basidia clavate, ca 5.5-6.5 um wide.
Sterigmata 4, ellipsoid becoming mitriform to fusiform. Spores narrow,
depressed or somewhat sinuous, 9.0-20.0(-26.5) x 2.5-3.0(-3.5) um from
print (Q = 3.0-9.0).

Specimen examined: on discarded ear of Zea mays on woodland path,
Binifaldo, Majorca, 10 Nov. 1992, P. Roberts 565, holotype, K(M) 28705

Though based on a single collection, the narrow, elongated spores of 7.
balearica are distinctive and unlike any other TJu/asnel//la species yet
described. 7. pa/lidocremea Julich has equally narrow but much shorter
spores (ca 10-12 um long) and _ substantially larger basidia. 7.
deliquescens (Juel) Juel also has narrow spores, but they are consistently
longer (ca 25-50 um) and of a different, vermiform shape. 7. ca/ospora
(Boudier) Bourdot & Galzin and 7. brinkmanni Bres. have spores of
approximately the right length, but fusiform and substantially wider (at
least 3.5-5.0 um across).

The type specimen's unusual substrate (a discarded ear of maize
found on a damp woodland path) is unlikely to be of consequence, since
most if not all 7u/asne//la species appear to be litter saprotrophs without
host preferences.

Additional species collected

All the following collections are from Majorca. The Kew herbarium number
is given where specimens have been retained.

Dacrymycetales

Calocera cornea (Batsch: Fr.) Fr. on fallen twig, Bosc de Monnaber, 13
Nov.1992, P.C. Holland

-Dacrymyces enatus (Berk. & M.A. Curtis) Massee, on fallen ?Quercus
branch, Bosc de Monnaber, 13 Nov. 1992, C.S.V. Yeates, K(M)
26215

Dacrymyces stillatus Nees: Fr., on fallen branch of Quercus cf. i/ex,
Menut, 10 Nov. 1992, P. Roberts; same substrate, Bosc de
Monnaber, 13 Nov. 1992, P. Roberts

123
Platygloeales

Helicogloea lagerheimii Pat., on fallen wood, Bosc de Monnaber, 13 Nov.
1992, P. Roberts 590, K(M) 26196

Tremellales

Eichleriella deglubens (Berk. & Broome) D.A. Reid, on partly fallen branch,
Binifaldo, 10 Nov. 1992, P. Roberts 572, K(M) 29745

Exidiopsis effusa (Bref. ex Sacc.) Maller, on fallen wood, Binifaldo, 10
Nov. 1992, P. Roberts 570, K(M) 26809

Myxarium nucleatum Wallr., on fallen wood, Menut, 10 Nov. 1992, P.
Roberts 557, K(M) 30617; on fallen branch, Binifaldo, 10 Nov.
1992, P. Roberts; on fallen twig, Formentor, 14 Nov. 1992, P.
Roberts.

Tremella foliacea Pers.: Fr. on fallen branch of Quercus cf. i/ex, Bosc de
Monnaber, 13 Nov.1992, A. Henrici, K(M) 29999

Tremella mesenterica Retz.: Fr., on fallen branch of Quercus cf. i/ex, Bosc
de Monnaber, 13 Nov.1992, P. Roberts

Tulasnellales

Tulasnella pallida Bres., Bosc de Monnaber, 13 Nov. 1992, P. Roberts
585, K(M) 26146

Tulasnella violea (Quélet) Bourdot & Galzin , Bosc de Monnaber, 13 Nov.
1992, P. Roberts 588, K(M) 26154

Thanks to Dr J. Ginns for kindly reviewing this paper.

REFERENCES

Oberwinkler, F. (1963) Niedere Basidiomyceten aus Stdbayern Ill. Ber.
bayer. bot. Ges. 36: 41-55

Roberts, P. (1993) Exidiopsis species from Devon, including the new
segregate genera Ceratosebacina, Endoperplexa, Microsebacina,
and Serendipita. Mycol. Res. 97: 467-478

Warcup, J.H. & Talbot, P.H.B. (1967) Perfect states of Rhizoctonias
associated with orchids. New Phytologist 66: 631-641

Wells, K. & Raitviir, A. (1975) The species of Bourdotia & Basidiodendron
of the USSR. Mycologia 67: 904-922

=
7 he
ie PLS - i
vy : tee
y vie 4
in ¢
0 ' i
— & j j i Po)
j =
és . aa e |
ue i
“%
+ IP »
‘ < e j 4
4 " my ay '
4%
x ‘
i i)
i 5 '
\ '
y
} | bie: (a8. BR ae |
ie ; 5 ny Ae i .
, eG @ledowsh St CelRGl wakole sii
4 j '
in et
We wa . ‘ é Bal a re ¥, KF
ik fit
+0 ’ f hay 4 ah
3 | } é
+ ;
i
y Ra! 4
4 n LEME RON ya PAA eT et. Shere

a ee ae ef ; if
ob italia: ta ii J Ce (Gs
tonne aidi Lainie fio, eeannllig eM
, : : P Pus u;
! 4 | ‘in a pn re F iciagaur, © My Hj Th a

i agp ; 464 ea bunl hes ok oe A &
ae tit te seers terrae OB" Drigts
’ , ’ , , “iw es

put ae rae lig # fey bay 1 f eye idl \ wits, si MG

"seca Ngnaagll ho | nis ie cohen ea bia

MYCOTAXON

Volume LX, pp. 125-128 October-December 1996

A NEW SPECIES OF CALLISTOSPORIUM

Raymond M. Fatto Alan E. Bessette

1187 Millstone River Road Biology Department

Somerville, NJ 08876-4726 Utica College of Syracuse University
1600 Burrstone Road

Utica, NY 13502
SUMMARY

Collections of a species of Callistosporium did not match published
macroscopic descriptions of Callistosporium luteo-olivaceum. A new
species, Callistosporium purpureomarginatum is described.

INTRODUCTION

A collybioid fungus found repeatedly while conducting a fungal inventory at
Rutgers University's Hutcheson Memorial Forest, Somerset Co., NJ, was recorded as
Callistosporium luteo-olivaceum (Berk.& Curt.) Singer primarily because of its
microscopic features. Disturbingly, the macroscopic features and habitat did not conform
with the literature description. When this same taxon was found at the 1994 NAMA
Foray at Montreat, NC, the species name "rubescentifolia" was suggested. A literature
search disclosed that Collybia colera var. rubescentifolia (Pk.) Pk. had been synonymized
(Bigelow and Barr, 1966; Redhead, 1982) with Cailistosporium luteo-olivaceum and
Peck's (1886) original macroscopic description and habitat did not equate with that of the
taxon in question. Since published descriptions do not satisfactorily portray this taxon, it
is here described as a new species.

MATERIALS AND METHODS

Color terms and designations used in the following descriptions are cited from
Maerz & Paul (1930) when designated as M&P 4J6 or from the ISCC-NBS system
(Kelly & Judd, 1955) when of the form, number, modifier, hue, e.g., 15 moderate red,
otherwise they reflect the authors' color concept. Microscopic descriptions were made
from sections mounted in water, 3% KOH and Melzer's solution. Unless indicated
otherwise, the specimens examined from NJ are deposited in the herbarium of the New
Jersey Mycological Association at Rutgers University, New Brunswick, NJ.

TAXONOMY
Callistosporium purpureomarginatum Fatto & Bessette sp. nov.

Pileus 10-45 mm latus, primo convexus ad marginem incurvatum tum late
convexus-pacne planus, sacpe aliquantum umbiliciformis, margo excedens lamellas, glaber

126

si humidus, ab fusco rubello-purpureo usque ad brunescentem-violaceum, hygrophanus,
discus mox luteolus-tannus usque ad roseolum-tannum sed in margine retinens fuscus
brunescens-violaceus. Contextus tenuis, pallidus-luteolus-tannus. Odor indistinctus.
Sapor leviter amerus, cum sapore metallico post aliquantum temporis. Lamellae adnexae
vel emarginatae, confertae, roseolae vel luteae cum purpureo tinctu supra, marginatae
rubellae-purpureae. Stipes 10-40 mm longus, 1.5-4 mm crassus, glaber, concolorus cum
lamellis, basis cum tomento albo vel cremeo. Basidiosporae 5-7 x 3-4.5 mm, formae
grani mali vel ellipticae, glabrae, inamyloideae. Cheilocystisia filiformis.

Holotypus legit Fatto 8-8-95, a R. M. Fatto lectus, in ligniis quercus cariosis,
prope Hutcheson Memorial Forest, Rutgers University, NJ, USA. 8 August 1995. in herb.
New York Botanical Garden conservatus.

Pileus (10-)15-30(-45) mm broad, convex with an incurved margin when young,
expanding to nearly plane, often subumbilicate, margin exceeding lamellae; surface
glabrous when young and moist to slightly radially fibrillose when dry, dark reddish purple
to brownish violet, M&P 7H6 to 6K10 or 7J1 to 6G10 or approximately 16 dark red to
46 grayish reddish brown when young and moist, hygrophanous, disc soon drying to a
yellowish tan, M&P 12J4 to 13H6 or 90 grayish yellow, and then fading to a pinkish tan,
M&P 4A9 to 12D5 or approximately 76 light yellowish brown from the disc towards the
margin but retaining a dark brownish violet band at the margin, band usually sharply
delineated, 1-3 mm wide,. the portion of the margin exceeding lamellae very fragile,
becoming blackish violet, when dried uniformly M&P 7A10 or 61 grayish brown
exclusive of the blackish marginal band; context thin, to 1 mm, pale yellowish tan; odor
pleasant but not distinctive; taste slightly bitter with a metallic aftertaste.

Fig. 1. Basidiocarps of Callistosporium purpureomarginatum. (Fatto) x 1.5

127

Lamellae adnexed to emarginate with a slightly decurrent tooth, of varying
lengths, close, thin, 1.5-3 mm broad, pinkish or yellow with purple overtones, edges
uneven, marginate reddish purple.

Stipe 10-40 mm long, 1.5-4 mm thick, equal, usually curved, occasionally
eccentric, hollow; surface glabrous to slightly fibrillose mostly on the upper half, slightly
striate, when fresh and moist concolorous with the lamella margins, hygrophanous, soon
purplish brown to yellow, aging brownish, base with white to creamy tomentum.

Chemical reactions: 3% KOH on pileus quickly medium to dark brown. 10%
NH,OH on pileus no reaction to faint browning.

Spore deposit white. Spores 5-7 x 3-4.5 ym, pip-shaped to elliptical, smooth,
inamyloid, hyaline, but in KOH, many uniguttulate with gray contents, in aged herbarium
spores the globule dissipates leaving grayed particles throughout the spore. Basidia 17-24
x 6-8 pum, clavate, four sterigmate. Pleurocystidia absent. Cheilocystidia filamentous,
24-40 x 2-4 um, cylindric to flexuous, apices rounded, some slightly capitellate. Lamellar
trama hyphae subparallel, 3-10 um wide. Pileipellis a layer of interwoven hyphae,
cylindrical, 3-10 tum wide, with protruding end cells, 30-50 x 5-8 um, 2-3 septate, with
fine encrusting particles. ileal trama hyphae interwoven, radial in orientation,
cylindrical to slightly inflated, 3-12 jum wide. Stipitipellis a layer of vertically oriented,
subparallel hyphae, 4-10 um wide. Clamp connections absent in ail tissues. Trama and
hymenium of fresh material pale amber, pileipellis and marginate lamella edges vinaceous
brown, and pileus margin strong vinaceous in water mount and remaining so, however, in
KOH trama and hymenium bright greenish yellow, pileipellis olive to pale vinaceous
brown, but all slowly changing to pale vinaceous, best seen when reexamined after 12
hours. All tissue of aged herbarium specimens as revived in KOH is immediately pale
brownish purple to vinaceous with dark purplish gray pigment inclusions or granules,
some internal, others encrusting.

Fruiting: Scattered to gregarious on well decayed oak logs; fruiting from mid July
to mid September; known from New Jersey and North Carolina, the exact range is
unknown.

MATERIAL EXAMINED: NEW JERSEY: Middlesex Co.: Rutgers University’s Helyar Woods,
15 August 1994, E. H. Vamey. Somerset Co.: Rutgers University’s Hutcheson Memonial Forest, 27 August
1991, Fatto; 7 August 1992, Fatto; 31 August 1993, Fatto; 12 July 1994, Fatto; 1 August 1995, Fatto; 8

August 1995, Fatto, (holotype of Callistosporium purpureomarginatum Fatto & Bessette), (NY). NORTH
CAROLINA: Buncombe Co.: Montreat, 4 September 1994, Bessette 9962 (NYS).

Observations: Callistosporium purpureomarginatum Is easily distinguished by the
reddish purple pileus, reddish purple marginate lamellae, the thin violet band at the pileus
margin, the reddish purple stipe which quickly dries to yellow and brown and is usually
curved because of growing out of the lower sides of decaying oak logs, and its collybioid
stature. Callistosporium luteo-olivaceum is typically yellowish brown overall, with gill
edges that are only occasionally reddish brown in age and grows on decaying conifer
wood. Baeospora myriadophylia has crowded, thin, lavender gills.

Once plucked from the cool, dark, high humidity environment in which C.
purpureomarginatum grows, the color change from reddish purple to yellowish tan
occurs rapidly, usually in 2 to 3 hours. The distinguishing features then are the grayish
yellowish tan pileus with a sharply delineated deep brownish violet band at the margin,
pinkish to yellow lamellae with a purple margin, and a yellow-brown stipe. KOH applied
to the pileus at this stage produces a deep brown color as dark as the band at the margin,

128

whereas if applied while the specimen is fresh and moist and still reddish purple, the color
merely becomes slightly deeper and browner. This brown color leaches out in a KOH
mount leaving the tissue a bright greenish yellow. At both these color stages, 10%
ammonium hydroxide has little effect; however, applied a day later, it produces a medium
brown color. This is truly a chameleon mushroom.

ACKNOWLEDGMENTS

We thank Dr. Roy Halling of NY and Dr. John Haines of NYS for their
assistance, and Alice Janick for providing the Latin description.

LITERATURE CITED

Bigelow, H. E. and M. E. Barr. 1966. Contribution to the fungus flora of northeastern
North America. IV. Rhodora 68: 175-191.

Kelly, K. L. and D. B. Judd. 1955. The ISCC-NBS Method of Designating Colors and
a Dictionary of Color Names. ISCC-NBS Color-Name Charts Illustrated with
Centroid Colors. National Bureau of Standards Circular 553. U.S. Government
Printing Office, Washington, DC. 158 pp.

Maerz, A. and M. R. Paul. 1930. A Dictionary of Color. McGraw-Hill Book Co., Inc.
New York, NY. 207 pp.

Peck, C. H. 1886. Report of the state botanist. Annual Report N.Y. State Museum of
Natural History 39: 30-75.

Redhead, S.A. 1982. The systematics of Callistosporium luteo-olivaceum. Sydowia
35: 223-235.

MYCOTAXON

Volume LX, pp. 129-136 October-December 1996

SCUTELLOSPORA CERRADENSIS: AN ORNAMENTED SPECIES
IN THE GIGASPORACEAE (GLOMALES)
FROM THE CERRADO REGION OF BRAZIL

JOYCE LANCE SPAIN

3899 NW Jameson Dr.
Corvallis, OR 97330

JEANNE CLAESSEN DE MIRANDA

CPAC/EMBRAPA CAIXA POSTAL 08223
PLANALTINA, D.F. BRAZIL 73301-970

ABSTRACT

A new species of arbuscular mycorrhizal fungi was found in
association with plants growing in a dark red latosol in a savanna
ecosystem. Pot cultures were established and observations made
of papillose hyaline to sub-hyaline spores mounted in polyvinyl
alcohol-lactic acid-glycerin (PVLG), water and Melzer's reagent.
Scutellospora cerradensis was characterized by a four layered
spore wall with a pigmented germination shield forming between two
flexible inner walls.

RESUMO

Uma espécie nova de fungos micorrizicos arbusculares foi
encontrada em associa¢cao com plantas cultivadas em um Latossolo
Vermelho Escuro em um ecossistema do tipo savana. Culturas em
vasos foram conduzidas e observacdédes foram feitas em esporos
papilosos e hialinos a sub-hialinos, montados em polivinil-alcool-
acido latico-glicerina (PVLG), agua e reagente de Melzer.
Scutellospora cerradensis foi caracterizada através da parede do
esporo, constituida de quatro camadas, com um escudo de germinacado
pigmentado formado entre duas flexiveis paredes internas.

INTRODUCTION

Scutellospora cerradensis, an early accession (1984) in the
collection of native arbuscular endomycorrhizae maintained at the
Centro de Pesquisa Agropecuaria dos Cerrados (CPAC), was initially
identified as S. verrucosa, a spore with similar ornamentation,
size and color. A more thorough study revealed a unique spore
distinguishable from S. verrucosa and other described species.

130

An innermost flexible layer of the spore wall, not previously
described, is discussed. Wall descriptions are based on
ontogenetic findings in the study of two Scutellospora species by
Franke and Morton (1994). A murograph depicts wall morphology.

MATERIALS AND METHODS

Spores, recovered by wet sieving soil and roots from a pot
culture, were suspended in water and cleaned by ultrasound for 30
seconds. Color determinations were made by stereomicroscope using
fiber optic illumination, color temperature 3200K. Intact spores
suspended in water against a black background were compared to the
INVAM Color Chart (1993). Color of shields, sporophores,
auxillary cells amd wall layer reactions to Melzer's reagent was
assessed against a white background.

The spore wall and inner walls of numerous spores were separated
in water or PVLG on a slide using a scalpel and dissecting
needles. Intact spores, ruptured spores and dissected spores were
mounted in PVLG, water or Melzer's reagent and examined with a
compound microscope. Melzer's reagent was also diluted with PVLG
or distilled water to observe the rate and intensity of
histochemical responses. The species diagnosis was based on
observations of ruptured and/or dissected spores with shields
mounted in PVLG and magnified 1000 X. Type specimens were mounted
in PVLG; unmounted spores were preserved in 5% formalin.
Observations and measurements were also made of ruptured spores
in water.

LATIN DIAGNOSIS
Scutellospora cerradensis Spain & Miranda sp. nov. (Figs. 1-3)

Sporae singulae in solo feruntur in apice aut circa apicem
cellulae sporogenae; raro autem sporae duae feruntur a latere unis
cellulae sporogenae. Sporae maturae hyalinae vel subhyalinae;
senescentes, stramineae (0/0 10/0) vel pallide luteobrunneae (0/10
40/0), globosae vel obovoideum (167-) 230-363 pm X (167-) 230-363
pm, raro irregulares. In PVLG sporae tunica constituitur stratis
quatuor: exteriore papilloso <1.0-1.0 pm crasso et adherente in
secundo strato rigido, 1.5-2.0 pm crasso et adherente in tertio,
subtiliter lamellato, 2-3 pm crasso; quarto membranaceo <0.5-1.0
pm crasso. In PVLG stratum papillosum et stratum secundum
disjunguntur. Stratum quartum et stratum tertium disjunguntur,
excepta conjunctione cellulae sporogenae. Murus internus primus:
stratum primum 1.0 pm crassum adheret secundo 3-5 pm crasso.
Murus internus secundus constituitur stratis tribus: primum <0.5-
0.5 pm crassum adheret strato secundo 4-6 (-10) pm crasso, sed
crassities varietur in PVLG. Stratum tertium, in quo secundum
adheret, 0.5-1.0 pm crassum, et raro cum papillis rotundis <1-1.5
pm diametro. Scutellum inter murum primum et secundum formatur.

131

SPORES are hyaline to sub-hyaline when mature, straw (0/010/0) to
a pale yellow-brown (0/10/40/0) with age; globose to ovoid, (167-)
230-363 pm X (167-) 230-363 pm, rarely irregularly shaped. Usually
spores are borne singly in soil at or near apex of sporogenous
cell, however, two spores are occasionally borne laterally from
one sporogenous cell. Spore contents, opaque white in immature
spores, become hyaline with maturity.

The spore wall consists of a weakly laminate layer surrounded by
two outer layers and an inner flexible layer, generally separate
from the other layers except at point of attachment to sporogenous
cell. Unless observed immediately the papillose layer, <1-1 pm
thick, papillae 1- 1.5 pm diam. to slightly oblong, up to 1.5 pm
high, will be perceived in PVLG as a fine reticulum or simply as
a roughened layer which separates from the rigid adjacent layer,
1.5-2.0 pm thick, adherent to laminate layer, 2-3(-5) pm thick.
The innermost flexible layer, <0.5-1.0 pm thick, is thicker at
connection to sporogenous cell. Inner wall one has an outer
layer, 1 pm thick, loosely adherent to an inner layer 3-5 pm
thick. Inner wall two has a very thin outer layer, <0.5 pm thick,
adherent to a thick layer, 4-6 (-10) pm, over a thin, sometimes
papillose (Fig. 3G), layer, 0.5-1.0 pm thick. The germination
shield forms between inner walls 1 and 2.

The ornamentation on the outermost layer of the spore wall is
discrete in water (Fig. 3A). The spore wall is of four layers:
together layer one, with papillae <1-1 pm thick with round, 1- 1.5
pm diam., to slightly oblong up to 1.5 pm high, and layer two are
approximately 1 pm thick; layer three, laminated, 2-3 pm thick;
layer four, the innermost layer, 0.5-1.0 pm thick, is rarely
observed. The outer layer of inner wall one, 0.5-1 pm thick,
adheres to the inner layer, 2-3 pm thick. Inner wall two has
three adherent layers: a middle layer, 2-4 pm thick, surrounded
by an outer layer, 1-1.5 pm thick, and an innermost layer, <1 pm
thick, observed as wrinkled or when sparse to more dense papillae,
<1-1.5 pm diam and 1-2 pm high, are present (Fig. 3G).

5 Phy eee 4 1 2 pia?
O
5S W IW-1 IW-2 IW-1 IW-2
PVLG Water
Figure 1. Murograph of Scutellospora cerradensis. Spore wall

(SW); Inner Wall One (IW-1); Inner Wall Two(IW-2). Open: rigid
Single layer; vertical dashes: laminae; diagonal lines: flexible
layer; hemispheres: plastic; 0: ornamented layer.

132

Figure 2. Scutellospora cerradensis germination shields, plan
view; camera lucida drawings. Aperture (A); ; Germ Tube Initial
(GTI); Fissure (F); ‘Y' configuration around aperture (Y).

B53

REACTION TO MELZER'S REAGENT. Five layers can be observed to
react with PVLG/Melzer's reagent (1:1). The laminated layer of the
spore wall is reddish yellow (INVAM 0/20 20/0) to reddish brown
(20/80 60/0) and clearly distinguished from the outer two layers;
the outer layer of inner wall one reacts very weakly and has a
tinge of pink (INVAM 0/20 20/0) when folded on itself; the inner
layer is pink (INVAM 0/60 20/0). Two layers of inner wall two are
observed to react: a light pink (INVAM 0/60 20/0) outer layer over
a reddish purple (INVAM 20/80 20/0) layer. In PVLG/Melzer's
reagent (3:1) the innermost layer (masked by the reaction of the
middle layer at a 1:1 ratio), appears to have a weak dextrinoid
reaction. A differential reaction sometimes observed in the
middle layer suggests that it may be, in fact, two layers.

GERMINATION SHIELD, light yellowish-brown to brown (INVAM 0/40
80/40 to 0/40 100/00), (87-)125-155 pm X (100-) 137.5-172.5 pm,
forms between inner walls one and two; a 'V' or ‘Y' configuration
develops around aperture (Fig. 7). The largest number of
germination tube loci observed was 6; the germination tube widens
from a diam. of 5 pm at egress to 12.5 pm near spore.

SPOROGENOUS CELL: light yellowish brown to brown, (INVAM 0/40 80/0
to 0/40 100/0) 35-55pm X (-30)50-75pm (width by distance between

apex and first septum of sporophore), often with fine hyaline
hyphae coming from one to several peg-like, pigmented hyphae.

Papillae are often observed on the innermost layer. A single spore
usually develops at apex, however, two spores, generally with a
disparity in size, occasionally emerge laterally from a single
sporogenous cell. The sporophore is septate and rarely branched
(Spain et al., 1989).

AUXILIARY CELLS 20-33 pm diam, smooth, round to knobby, formed in
soil on yellowish brown to brown (INVAM 0/40 80/0 to 0/40 100/0)
coiled or straight hyphae 3.5-4 pm diam, wall <ipm thick.

MYCORRHIZAL ASSOCIATIONS. Known to associate and develop
arbuscules with Allium porrum, Stylosanthes guyanensis var.
vulgaris cv Mineirado and Sorghum bicolor L. Moench, host plants
for pot culture.

DISTRIBUTION AND HABITAT: Spore known from experimental fields at
CPAC/EMBRAPA, Planaltina, D.F. Brazil; 15°935'33" south; 47942'30"
west; 1100 m altitude. This accession was isolated from a dark
red latisol (mostly clayey) with poor phosphorus availability.
Soil analyses: pH (water 1:2.5):4.8-5.0; phosphorus (ug/g soil):
3.5; aluminum (me/100 g soil): 0.2; calcium + magnesium (me/100
g soil)3.5; potassium (ug/g soil):40. Vegetation at the
collection site progressed from predominantly native grasses to
pasture to soybeans. A fallow period of about five years was
followed by a planting of different legumes for green manure. The
isolate was recovered from plots cultivated with Tephrosia
candida, however, an association with that plant has not been
determined.

134

Figure 3. Scutellospora cerradensis. A. Spore Wall layer 1
papillose, water, 1000X. B. Spore Wall, layer one detached, PVLG,
200X. C. Spore wall, layer one missing, PVLG, 400X DIC. D. Inner
Wall 1, two layers; Inner Wall 2, three layers; Spore Wall layer
four visible above Inner Wall 1, PVLG 400X. E. Inner Wall 1, two
layers, layer one wrinkled; both reactive to Melzer's reagent,
200X DIC. F. Inner Wall 2, three layers, layer 2 very plastic,
PVLG, 200X DIC. G. Inner Wall 2, papillae on layer three rare.
PVLG, 400X. H. Germination shield, PVLG 400X.

135

ETYMOLOGY: Place: cerrado; ensis. Epithet chosen to associate the
species with the savannah ecosystem (cerrado) where it was found.

TYPE: From endomycorrhizal association with Sorghum ‘bicolor L
Moench in pot culture Scutellospora sp. CPAC-3 at CPAC/EMBRAPA,
Planaltina, D.F. Brazil. Holotype: OSC; isotypes FH, K and IBt.
Cultotype, isolate BR 103, deposited at INVAM (West Virginia
University, Morgantown, WV).

DISCUSSION

Scutellospora cerradensis and S. verrucosa (Koske & Walker) Walker
& Sanders have similar ornamentation, size and somewhat similar
color when viewed in water against a white background by
stereomicroscope, however, the shield is patent only in S.
cerradensis. Scutellospora verrucosa, member of a sub-group
including S. coralloidea, S. fulgida, S. gregaria and S. persica
(Morton, 1995), lacks the second inner wall found in S.
cerradensis. Mounted spores of § S. cerradensis lacking
ornamentation might also be confused with S. gilmorei (Trappe &
Gerd.) Walker & Sanders, a smooth spore of similar size, color and
wall number.

The spore wall of S. cerradensis has four phenotypically distinct
layers. The outer layer is unique in its reaction to PVLG; unlike
other described ornamented spores, the papillae are not
structurally stable (3A). The ornamented layer reacts almost
immediately and can appear as a fine reticulum or a roughened
surface before detaching from the adjacent smooth layer (3B).
Layer three was rarely perceived as laminate; the very thin and
flexible innermost spore wall layer was not seen initially in
water mounts. Although often appearing in PVLG mounted spores to
be associated with the inner walls (3D), it clearly originates
from the sporogenous cell (3B & 3C). It is not known whether
there is discontinuity in the appearance of this layer as related
to the other spore wall layers. This very thin flexible layer,
not reported in the ontogenetic study of S. heterogama and S.
pellucida (Franke & Morton, 1994), is present in both species and
is not a species level character. Examination of a broader range
of taxa in Scutellospora will help determine if it is a sub-genus
or genus-level character.

The second inner wall is comprised of three layers (3D & 3F).
This wall is a highly conserved primary character (Franke &
Morton, 1994; Morton, 1995) and Scutellospora species described
as having a bi-layered second inner wall are hypothesized to be
equally complex. Papillae on the innermost layer of some spores
(3G) are not considered a taxonomic character. Numerous spores
of S. gregaria, an isolate native to the cerrado, develop papillae
on the innermost layer prior to shield formation (Spain, unpub.).
Papillae have also been observed on the innermost layer of an
isolate of S. verrucosa, a spore with one flexible inner wall.
It is not known whether this novelty is genetically induced or a
response to biotic or abiotic factors.

136

Different mountants affect morphology and histochemical reactions
of structures in spores of arbuscular fungi (Morton, 1990, Spain,
1990) The subcellular characters were detected and interpretable
only with detailed comparisons in both water and PVLG. Itsis
suggested that the example of Berch & Koske (1986) and Walker et
al. (1993) be followed: diagnoses should be made from observations
of spores mounted in water and PVLG. Although ontogenetic studies
were not carried out with S. cerradensis, most of the
developmental stages described by Franke & Morton (1994) were
observed. If found in other habitats S. cerradensis should be
readily identifiable.

ACKNOWLEDGEMENTS

We wish to thank Valter Lopes, long time assistant of Dr. Miranda
at CPAC/EMBRAPA. We are grateful to Dr. Chris Walker for valuable
suggestions and Dr. Joseph Morton for the critical review of the
manuscript. Father Hugh Feiss OSB kindly provided the Latin
translation and Dr. Stephen Alderman assisted with DIC

photography. The senior author acknowledges the cooperation
kindly given at CPAC/EMBRAPA in Brazil and the lab of Dr. Gabor
Bethlenfelvay in Corvallis. Dr. James M. Spain was a valued
critic.

LITERATURE CITED

Berch, S.M. and R.E.Koske. 1986. Glomus pansihalos: a new species
in the Endogonaceae, Zygomycetes. Mycol. 78:838-842.

Franke, M. and J. Morton, 1994. Ontogenetic comparisons of
arbuscular mycorrhizal fungi Scutellospora heterogama and
Scutellospora pellucida: revision of taxonomic character
concepts, species descriptions, and phylogenetic hypotheses.
Can. J. Bot. 72:122-134.

Gerdemann, J.W. & J.M. Trappe. 1974. The Endogonaceae in the
Pacific Northwest. Mycologia Memoir No. 5:1-76.

Morton, J.B. 1995. Taxonomic and phylogenetic divergence among
five Scutellospora species based on comparative developmental
sequences. Mycol. 87(1):127-137.

Morton, J.B. 1987. Effects of mountants and fixatives on wall
structure and Melzer's reaction in spores of two Acaulospora
species (Endogonaceae). Mycologia 78:787-794.

Spain, J.L. 1990. Arguments for diagnoses based on unaltered wall
structures. Mycotaxon 38:71-76.

Walker, C., V. Gianinazzi-Pearson, and H. Marion-Espinasse, 1993.
Scutellospora castanea, a newly described arbuscular
mycorrhizal fungus. Cryptog. Mycol. 14:279-286.

MYCOTAXON

Volume LX, pp. 137-142 October-December 1996

GLOMUS BRASILIANUM: AN ORNAMENTED SPECIES
IN THE GLOMACEAE

JOYCE LANCE SPAIN

3899 NW Jameson Dr. Corvallis, OR 97330

JEANNE CLAESSEN DE MIRANDA

EMBRAPA/CPAC Caixa Postal 08223
Planaltina, D.F. Brazil 73301-970

ABSTRACT

Glomus brasilianum, a hyaline, vesicular-arbuscular species with
labyrinthiform ornamentation on an inner layer, was recovered from
a dark red latosol in the experimental area of EMBRAPA/CPAC
(Centro de Pesquisa Agropecudria dos Cerrados), Planaltina, D.F.
Brazil. The description is based on a study of recently harvested
spores mounted in water and in polyvinyl alcohol-lactic acid
glycerol (PVLG), a mountant which induced morphological change.

RESUMO

Glomus brasilianum, uma espécie hialina, vesiculo-arbuscular com
ornamenta¢ao labirintiforme em uma camada interna, foi coletada
de um Latossolo Vermehlo Escuro na Area experimental da
EMBRAPA/CPAC (Centro de Pesquisa Agropecudria dos Cerrados),
Planaltina, D.F. Brasil. A descric&o é baseada em um estudo com
esporos coletados recentemente e montados em Agua e em polivinil-
alcool-acido latico glicerol (PVLG), um montante indutor de
modifica¢gées morfoldégicas.

INTRODUCTION

The native spore collection of vesicular-arbuscular mycorrhizae
at the Cerrado Center includes several unidentified Glomus species
in addition to G. convolutum Gerdemann & Trappe, G. manihotis
Sieverding & Schenck and G. occultum Walker. Glomus brasilianum,
initially found as a contaminant in a pot culture, was later
recovered in field sampling. A detailed study revealed unique
morphology.

138

Figure 1. Glomus brasilianum A) Spore wall, water i000X.
B) Spore wall, PVLG 1000X. C) Spore wall, Melzer's reagent 1000X.
D) Labyrinthiform ornamentation, layer 3, water 1000X. E)
Sporophore ornamentation, water 1000X. F) Plug, water 1000X G)
Germination through spore wall, water 400X. H) Halo effect;
intact spore, PVLG 400X. I) Knobby intercellular hypha 1000X.

MATERIALS AND METHODS 139

Spores isolated from a contaminated pot culture were inoculated
into pots using Allium porrum as the host plant. A pure culture
was obtained and spores, recovered by wet sieving with a fine mesh
sieve (45pm), were examined by stereomicroscope against white and
black backgrounds for color determination. The description is
based on observations of intact and ruptured spores mounted in
water, PVLG and Melzer's reagent examined at 1000X with a compound
microscope. Type specimens were permanently mounted in PVLG;
unmounted specimens were preserved in 5% formalin.

DESCRIPTION

Glomus brasilianum Spain et Miranda sp. nov. Figs. (1 & 2).

Sporocarpia ignota. Sporae, hyalinae, communiter globosae (47.5-)
60-80 (-112.5) pm vel ellipsoideae (47.5-) 64 X 77 (-115) um,
singulae in solo feruntur in sporophoribus et raro in radiis
senescentibus. Tunica sporae constituitur stratis quinque (ex
mensuris sporarum ruptarum in aquis): primo, mucilagino, 0.5-<1.0
ym crasso, communiter cum detritu affixo et in secundo adherente;

secundo, duro fragilique, 1 wm crasso; tertio, 1-2 wm crasso,
forma labyrinthi ornato, adherente in quarto quod est lamellatum

flexibileque, 1-3 wm crassum, adherens quinto, quod est
membranaceum, 0.5-<1.0 pm crasso. Tubae germinativae tunicam
penetrant.

Sporocarps unknown. Spores, hyaline, 2 SRS

generally globose (47.5-) 60-80 (-112.5) um
or ellipsoid, (47.5-) 64 X 77 (-115) um,
borne singly on sporophore in soil and
occasionally in senescent roots. Spore wall
structure of five layers (measurements of
ruptured spores in water). Spore wall layer
1, mucilagenous, 0.5-<1 pm thick usually
with debris attached, generally adherent to
layer 2, a brittle layer to ipm thick; layer
3 with labyrinthiform ornamentation, 1-2 pm

1

Sl
y
;
f
;
,

thick, adherent to layer 4, a flexible ra)
laminated layer 1-3 pm thick, greatest

thickness near sporophore; layer 5, Figure 2.
membranous, 0.5-<1 wpm thick, usually Murograph of
appressed to layer 4, occasionally spore wall
separating, more often discerned by structure,
wrinkling (Fig. 1B). The germ tube emerges water mount.

through the spore wall.

REACTION TO COMMON MOUNTANT PVLG AND MELZER'S REAGENT: In PVLG,
spore wall layer 1, mucilagenous, <1l-ipm thick, often adheres,
partially adheres or separates from layer 2, 1-2 pm thick. Layers
2 and 3 separate as layer 3, the ornamented layer, 1-2 pm thick
initially, swells, losing definition. Layer 3 adheres to flexible
laminated layer 4, 1-2.5 wm, laminae swelling and separating
slightly at point of rupture; layer 5, membranous, 0.5 pm thick
often separates from layer 4 (Fig. 1B). Over time a halo effect

140

is created in many intact spores as the ornamented layer swells

(Figs: 1-B)k The dextrinoid reaction is absent in Melzer's
reagent, however, the walls absorb pigment becoming yellow even
in a dilute solution. The ornamented layer usually swells

markedly, to 5 pm, creating striations (Fig. 1C).

GERMINATION: Germination tube emerges through the spore wall
(Fig. 1G).

SPOROPHORE: Usually straight, sometimes curved, 5-6pm in
diameter, generally coenocytic, often narrowing slightly near
spore. Layers of sporophore wall are contiguous with the three
outer layers, however, layers 1 and 2 are often missing in sieved
spores and the ornamented layer is outermost (Fig. 1E).

MYCORRHIZAL ASSOCIATIONS: Glomus brasilianum is known to form
mycorrhizae with Allium porrum, Sorghum bicolor L. Moench and
Stylosanthes guyanensis var. vulgaris cv Minerao, host plant used
for pot cultures. The infection is characterized by intracellular
coiled hyphae and intercellular hyphae; projections are sometimes
present on both (Fig. 1-I). Vesicles mounted in PVLG after
staining with trypan blue are generally distinguished by a 2 pm
thick wall; arbuscules have extremely fine ramifications and are
observed as a very lightly stained mass when senescent.

DISTRIBUTION AND HABITAT: Glomus brasilianum, known first as a
contaminant in the greenhouse (potting soil has a pH of 4.8 (2.5:1
HOH); Al 2.11 me/100g; Ca + Mg: 0.16 me/100g; P: 0.97 ppm; K: 12
ppm.) was also found in an experimental pasture planted to
Brachiaria decumbens (dark red latosol with 4-5 ppm available P)
at EMBRAPA/CPAC, Planaltina, D.F. Brazil (15° 35'33" south; 47°
42°30"; 1100 m altitude).

ETYMOLOGY: brasilianum, place Brazil, indicating that the species
was first observed in Brazil.

TYPE: Spores recovered from pure pot culture Glomus sp. CPAC-5,
an association of G. brasilianum with A. porrum. Type OSC;
Isotypes: FH, K and IBt, INVAM Accession: BR105.

DISCUSSION

Morphological changes caused by mountants and preservatives were
studied by Morton (1986, 1988) and noted by others (Berch &
Fortin, 1983; Schenck et al., 1984; Berch & Koske, 1986). It has
been suggested that descriptions of some spores be made from water
mounts because altered walls do not represent spore morphology
(Spain 1990). Spores of G. brasilianum are dramatically altered
by both PVLG and Melzer's reagent. The ability to distinguish the
detail of the ornamentation on spores mounted in PVLG or Melzer's
reagent seems to be a function of time rather than spore age; the
nature of the ornamentation generally is only apparent if
observations are made in water or immediately following slide
preparation when spores are mounted in PVLG.

141

Layers 1 and 2 are often entirely missing from sieved spores and
the ornamented layer, contiguous with the sporophore (Fig. 1E),
appears as the outer wall. The flexible laminated layer, first
described by Koske and Gemma (1989), is rarely observed as
laminate and perusal of numerous spores may be necessary to
corroberate the finding. The layer is thicker near the sporophore
(Fig. 1E & F) and usually has a slightly greater thickness at
rupture sites. When appressed to layer 4, layer 5 is discerned
because of wrinkling; the spore contents seem to be isolated by
a plug (Fig. 1F).

Several observations were made of spores with a single germination
tube emerging through the spore wall (Fig. 1G); occasionally the
germination tube and the sporophore hypha anastomosed. Germinated
spores with detached sporophore and germination tube, essentially
empty of spore contents, also exhibited two openings in the spore
wall. Glomus albidum (Walker and Rhodes, 1981) and G. pallidum
(Hall, 1977) are also reported to germinate directly through the
wall.

Different types of multiple walls (layers) account for most of the
morphological diversity in the Glomaceae (Morton & Benny, 1990).
Glomus brasilianum is an anomaly among described species having
an ornamented inner structural layer. It is easily distinguished
from G. maculosum Miller & Walker, a much larger spore with
bulging discs not found in hyaline immature spores but present on
the innermost membranous wall in many older straw-colored to
ochraceous spores (suggesting a possible relationship to
germination). Other hyaline spores are reported to have
ornamentation on the outer layer: G. callosum Sieverding is
ornamented with crowded minute warts; with age the mucilaginous
coat of G. clarum Nicolson & Schenck becomes verrucose or rugose;
G. leptotichum Schenck & Smith has an indistinct alveolate
reticulum most apparent on young spores; G. scintillans Rose &
Trappe is ornamented with hyaline knobs. The ornamentation of G.
brasilianum resembles that of Acaulospora rehmii Sieverding and
Toro (1987), however, it is unlikely that these species of
different genera would ever be confused. Although the sporophore
of G. brasilianum may detach, the very delicate labyrinthiform
ornamentation on an inner structural layer distinguishes it from
sessile spores of A. rehmii with similar, but much more prominent,
ornamentation on the spore wall.

ACKNOWLEDGEMENTS

The authors wish to acknowledge the support of CPAC/EMBRAPA. We
are grateful for the assistance of Valter Lopes and José Moreis
in the laboratory. We are indebted to Chris Walker for reviewing
the manuscript and making helpful suggestions. Father Hugh Feiss
OSB kindly provided the Latin diagnosis. The senior author wishes
to thank James M. Spain for his encouragement and valuable
criticism.

142

LITERATURE CITED

Berch, S.M. and J.A. Fortin, 1983. Lectotypification of Glomus
macrocarpum and proposal of new combinations: Glomus australe,
Glomus versiforme, and Glomus tenebrosum (Endogonaceae). Can.
J. of Bot. 61:2608-2617.

Berch, S.M. and R.E. Koske, 1986. Glomus pansihalos, a new
species in the Endogonaceae, Zygomycetes. Mycologia. 78:832-
836.

Hall, I. 1977. Species and mycorrhizal infections of New Zealand
Endogonaceae. Trans. Br. Mycol. Soc. 68:341-356.

Koske, R.E. and J.N. Gemma 1989. Glomus nanolumen (Endogonaceae),
a new species from Hawaii. Mycologia 81935-9387

Morton, J.B. 1986. Effects of mountants and fixatives on wall
structure and Melzer's reaction in spores of two Acaulospora
species (Endogonaceae). Mycologia 78:787-794.

Morton, J.B. 1988. Taxonomy of VA mycorrhizal fungi:
classification, nomenclature, and identification. Mycotaxon
32: 267-324.

Morton, J.B. and G.L. Benny 1990. Revised classification of
arbuscular mycorrhizal fungi (Zygomycetes): A new order,
Glomales, two new suborders, Glomineae and Gigasporineae, and
two new families, Acaulosporaceae and Gigasporaceae, with an
emendation of Glomaceae. Mycotaxon 37:471-491.

Schenck, N.C., J.L. Spain, E. Sieverding and R.H. Howeler, 1984.
Several new and unreported vesicular-arbuscular mycorrhizal
fungi (Endogonaceae) from Colombia. Mycologia 76:685- 699.

Sieverding, E. and S. Toro T. 1987. Acaulospora denticulata sp.
nov. and Acaulospora rehmii sp. nov. (Endogonaceae) with
ornamented spore walls. Angewandte Botanik 61:217-223.

Spain, J.L. 1990. Arguments for diagnoses based on
unaltered wall structures. Mycotaxon 38:71-76.

Walker, C. and L.H. Rhodes 1981. Glomys albidus: A new species in
the Endogonaceae. Mycotaxon: 12: 509-514.

MYCOTAXON

Volume LX, pp. 143-148 October-December 1996

ROYOPORUS - A NEW GENUS FOR FAVOLUS SPATHULATUS
ACB De
Department of Botany, Burdwan Raj College, Burdwan-713 104, W.B.,India
SUMMARY

Detailed observations on morphological and
anatomical characters of Favolus' spathulatus
(Jungh.)3) Lev, are given bdsed on ifres

basidiocarps from India. In the light of these
Observations =~ the. \new senus Royoporus,., Deis

proposed with Favolus' spathulatus (Jungh.)
Lev. as its type species.

During a mycological excursion fresh basidiocarps of a _ shortly
stipitate poroid fumgus of the Aphyllophorales (Polyporaceae) were
collected by the author from a branch of a dead angiosperm at
Amarkantak, Madhya Pradesh, India on October 20, 1993. After investi-
gations this fungus was identified as Favolus spathulatus (Jungh.)
Lev. The present paper reports the results of detailed studies on the
morphological and anatomical characters of F. spathulatus. The
taxonomy of this species is also discussed in the light of these
findings. The combination of features which characterizes this fungus
is not found in any other genus and, therefore, a new generic name is
proposed :

Royoporus De, nov. gen.

Carpophorus stipitatis, pileus spathulatus vel flabellatus, stipus
lateralis, contextus albus, hymenio albido vel ochraceo, poris
hexagonis, radialiter instructus. Systema hypharum dimiticum. Hyphae
generativae hyalinae, non-fibulatae. Basidia hyalina, clavata, 4-
sterigmatibus, 18.4-22.0 x 3.0-6.0 pm, basidiosporae 5.0-8.0 x 2.0-
3.0 pm, cylindricae, levis, hyalinae, tenuitunicatae, non-amyloideae.

SPECIES TYPICAE : Laschia spathulata Jungh. ,Fl.Crypt.Java,75-76,1838.
ETYMOLOGIA : Secandum nomen Dr. Anjali Roy.

Basidiocarp stipitate, pileus spathulate to flabelliform, stipe
-lateral, context white, up to 1 mmm thick, hymenium white to
ochraceous, pores hexagonal, radially (ean anais Hypal system dimitic
with hyaline, simple septate generative hyphae and hyaline binding
hyphae, Basidia hyaline, clavate, 4-sterigmate, 18.4-22.0x3.0-6.0 jum
Basidiospores 5.0-8.0 x 2.0-3.0 pm, cylindrical, smooth, hyaline,

thin-walled, non-amyloid.

144

145

Royoporus spathulatus (Jungh.) De, Comb. nov.

= Laschia spathulata Jungh., Fl. Crypt. Java, 75-76, 1838 (basionym) ;
Favolus spathulatus (Jungh.) Lev., Ann. Sci. nat. Ser III, bot.,
Paris, 2 : 203, 1844 ; Favolus moluccensis Mont., Ann. Sci. nat.
Ser. II. bot., Paris, 20 : 365, 1843 ; Polyporus moluccensis (Mont. )
Ryv., Mycotaxon, 38 : 84, 1990.

Basidiocarp annual, solitary or several growing from a common point on
the substratum, coriaceous and corky, very shortly stipitate ; pileus
spathulate, flabelliform or dimidiate, tapering towards the base,
upper surface (Fig.l) flat or slightly convex, may be depressed near
the stipe, straw coloured with sharp fine radial striations ; margin
thin, entire or lobed, sterile below ; hymenial surface (Fig.2) white
to ochraceous, pores hexogonal, radially aligned, 1-4 per mi,
dissepiments very thin, tubes up to 3 mm long ; context white, up to
1 mm thick. Stipe very short, lateral, about 5 mm in diameter and up
to 8 mm long.

Hyphal system dimitic. Generative hyphae (Fig.3) hyaline, thin-walled,
rarely slightly thick-walled, simple septate, branching not common,
1.5-3.5 jam wide, abundant in trama and rare in context. Binding hyphae
(Fig.4) hyaline, corrugated, thick-walled, in the context showing wide
lumina, in trama frequently subsolid to solid, usually dichotomously
branched, 3.0-8.2 pum wide, often tapering to 1.5-2.0 pm. Gloeoplerous
hyphae (Fig.5) present, up to 8 pm wide. Basidia (Fig.6) hyaline,
clavate, 4-sterigmate, 18.4-22.0 x 3.0-6.0 pm. Basidiospores (Fig.7)
hyaline, thin-walled, smooth, cylindrical, non-amyloid, 5.0-8.0 x
2.0-3.0 pum, some with one or two guttulae. Cystidioles (Fig.8) 18.0-
24.0 x 3.0-6.0 pum.

TYPE OF ROT : White rot.

SPECIMEN EXAMINED : BRCMH P-1993, Madhya Pradesh, India, on dead
branch of an unknown angiosperm, 20.10.1993. Kept in the Mycological
Herbarium, Department of Botany, Burdwan Raj College, Burdwan
West Bengal, India (BRCMH). The duplicate material, (HCIO 41026) has
been deposited in the Herbarium of the Division of Mycology and Plant
Pathology, Indian Agricultural Research Institute, New Delhi, India.

ETYMOLOGY : The generic epithet after the celebrated mycologist of
India Dr. Anjali Roy (Department of Botany, Visva-Bharati University,
Santiniketan, West Bengal, India).

DISCUSSION

The genus Favolus Fr. (Elench. Fung., P.44, 1828) was previously
regarded as distinct from the genus Polyporus Fr. (Syst. Myc. 1: 341,
1821) due to lack of clamps in the former and their presence in the
latter (Ryvarden and Johansen 1980). But Stankovicova (1973) in a
detailed study of the hyphal system of Favolus brasiliensis (Fr.) Fr.
[=Favolus tenuiculus Beauv.], the type species of the genus Favolus
Fr., described clamps. Gilbertson and Ryvarden (1987) and Ryvarden
(1990) agreed, and they corrected the error in Ryvarden and Johansen

Figs.1 & 2. Royoporus spathulatus (Jungh.) De : l.upper sutface of
basidiocarp ; 2. hymenial surface of basidiocarp.

146

147

(1980) which stated the hyphae were simple septate. Due to presence of
clamps in the type species of both of. these genera, Favolus could
not be kept as a separate genus and Corner (1984) proposed to merge
Favolus Fr. with Polyporus s. str. This view was supported by Ryvarden
(1990) and Teixeira C1502). Gilbertson and Ryvarden (1987), Roy De and
Pal (1994) and Teixeira (1994) have accepted the transfer of Favolus
tenuiculus Beauv. (= F. brasiliensis), the type species of the genus
Favolus Fr., to the genus Polyporus Fr. made by Fries (1821). But
Favolus spathulatus (Jungh.) Lev. having simple septate generative
hyphae (Ryvarden and Johansen 1980) could not be transferred to
Polyporus along with F. tenuiculus and generic position of F.
spathulatus became uncertain. In 1990 Ryvarden transferred F.
spathulatus to the genus Polyporus Fr. and he made a new combination
Polyporus moluccensis (Ryvarden 1990) although he did not explain
whether clamps were observced by him in this species. Mention may be
made in this connection that Junghuhn's name Laschia spathulata of
1838 is the oldest name for this taxon but when it was transferred to
the genus Polyporus, Ryvarden (1990) could not coin the name Polyporus
spathulatus as this name was preoccupied by another polypore
[Polyporus spathulatus (Hook.) Fr. Epior. Mycol. 1838]. Montagne
described this species in 1843 as Favolus moluccensis Mont. [Ann. Sci.
nat. Ser. II, bot., Paris, 20 : 365, 1843 | and his name was therefore
used by Ryvardon (1990). Thus Polyporus moluccensis is the name for
the fungus (F. spathulatus)in the genus Polyporus.

But my study based on fresh basidiocarps revealed that F. spathulatus
has simple septate generative hyphae. Thus its inclusion in Polyporus
Fr., a genus characterized by the presence of clamps in all species,
can not be agreed upon. Moreover, F. spathulatus exhibits a
combination of characters that can not be found in any other existing
genus : (a) spathulate to flabelliform basidiocarp with very short,
lateral stipe, (b) poroid hymenial surface with radially aligned
hexagonal pores, (c) white context, (d) dimitic hyphal system with
hyaline, simple septate generative hyphae and hyaline binding hyphae,
(e) hyaline, thin-walled, smooth, cylindrical, non-amyloid
basidiospores, and (f) an association with a white rot of angiospermic
wood.

I, therefore, propose a new genus Royoporus De to accommodate F.

spathulatus and the transfer F. spathulatus as : Royoporus spathulatus
(Jungh.) De, comb. nov. (Basionym : Laschia spathulata Jungh., FI.

Crypt. Java, 75-76, 1838).

ACKNOWLEDGEMENTS

The author is indebted to Dr. Leif Ryvarden, University of Oslo,
Norway for confirming the identification of the fungus and for his
valuable comments. The author is also indebted to Dr. J. Gimns,
Curator, National Mycological Herbarium of Canada, for critically
reviewing the manuscript.

Figs. 3-8. Microscopic structures from basidiocarp of Royoporus
spathulatus (Jungh.) De : 3. generative hyphae ; 4.
binding hyphae ; 5. gloeoplerous hyphae ; 6. basidia ;
7. basidiospores ; 8. cystidioles.

148

REFERENCES

Corner, E.J.H.1984. Ad Polyporaceas II & III. Beiheft. Nova Hedwigia,
78 3: 1-222.

Fries, E.M.- 1821. Systema mycologicum, Vol. I. Lumdae, P. Lvii + 520. .

Gilbertson, R. L. and L. Ryvarden, 1987. North American polypores,
Vol. 2, Fungiflora, Oslo, Norway.

Roy, Anjali, A.B. De and S. Pal. 1994. The genus Polyporus, s. str.
Fr. in andia. Crypt. Bot.) 4:6 73381-3935.

Ryvarden, L. 1990. Type studies in the Polyporaceae 22. Species
described by C. G. Lloyd in Polyporus. Mycotaxon, 38 : 83-102.

Ryvarden, L. and Johansen, 1980. A preliminary polypore flora of East
Africa. Fungiflora, Oslo, Norway.

Stankovicova, L. 1973. Hyphal structure in some pleurotoid species of
Agaricales. Nova Hedwigia, 24 : 61-120.

Teixeira, A. R. 1994. Genera of Polyporaceae : an objective approach.
Boletim da chacara Botanica de Itu. 1: 1-91.

MYCOTAXON

Volume LX, pp. 149-174 October-December 1996

CHYTRIDIOMYCETE TAXONOMY SINCE 1960

Joyce E. Longcore

Department of Plant Biology and Pathology
University of Maine
Orono, Maine 04469-5722 U.S.A.

ABSTRACT: More than 300 species and 23 genera of Chytridiomycetes have been described
since F.K. Sparrow's 1960 edition of Aquatic Phycomycetes, which contained monographic
coverage of the class. Furthermore, recent authors have presented many new combinations and
suggestions for synonymy. This bibliography is organized alphabetically by taxon and lists all new
taxa and taxonomic changes that have been published since 1960. Entries for new species include
information about substrates and habitats.

KEY WORDS: Blastocladiales, Chytridiales, Chytridiomycetes, Monoblepharidales,
Neocallimastigales, Spizellomycetales, taxonomy

INTRODUCTION: Thirty-five years ago F.K. Sparrow (1960) published a monograph that
covered the Chytridiomycetes and other aquatic, zoosporic fungi. This comprehensive work
brought together all taxonomic literature on chytrids (excluding the genera Synchytrium and
Physoderma) since the first report by Braun in 1856. Further contributions to chytrid systematics
include a key to genera (Sparrow 1973), and keys to some genera and species in a book on aquatic
mycology (Batko 1975). Karling's (1977) Chytridiomycetarum Iconographia represents a
significant contribution. He discussed the genera of Chytridiomycetes and presented many useful
drawings, but did not include a comprehensive list of species. More recently, Barr (1990)
reviewed current ideas of the phylogeny of the Chytridiomycetes, but no monograph has
superseded that of Sparrow. New species continue to be described and taxonomic changes
continue to be published, but articles are dispersed in many journals and searches to locate all
described species in a genus are time consuming. The /ndex of Fungi contains entries for most
new species and taxonomic changes. All classes of fungi are included in the Index and it is
arranged by date, thus, a search for all species in a particular genus requires an issue by issue
search of several volumes. [Volumes 1-4 of the Jndex are now available electronically from the
United States National Fungus Collections and can be searched by genus or species for 1940-1980.
(TELNET FUNGI.ARS-GRIN.GOV,; at “OK,” type: login user; at “Password?” type: user)]
The following bibliography, which includes new taxa, new combinations, and suggested
synonymies, differs substantially from the Index of Fungi because the list brings together,
alphabetically by genus, all chytridiomycete taxonomic references for 1960-1995. Without the
ability to easily search for all described species in a genus, identifications are difficult and earlier
homonyms are not detected. For example, Rhizophydium spinosum has been used as a name for a
new species by three authors since Sparrow (1960).

150

This compilation is only a listing and inclusion herein does not reflect my opinion of the status of
a species or change in classification. Spelling of names is that given in the Index of Fungi where,
in some instances, orthographic corrections were made. New species in certain parasitic genera
are not included, namely: Coelomomyces and Physoderma (Blastocladiales) and Synchytrium
(Chytridiales). Because Harpochytrium was not included in Sparrow's monograph, I have
included species described before 1960. Citations contain inclusive pagination for articles instead
of only the page containing the description or the taxonomic change. This variance from
convention is to facilitate the retrieval of entire articles via interlibrary photocopy services. Names
of new taxa at the genus level and above are printed entirely in upper case letters. Except where
noted, I have examined photocopies or reprints of all cited articles. Figures, substrates, and
habitats are listed only for new species. This list is maintained as a database file and will be
updated. Please send additions and corrections to the author.

ABBREVIATIONS AND DEFINITIONS: Blastocladiales, [B]; British Columbia, B.C.;
Chytridiales, [C] ;California, CA; Chytridiomycetarum Iconographia (Karling 1977), CI; county,
Co.; English Lake District, E.L.D.; Florida, FL; Handbook of Protoctista (Barr 1990), HP;
Hawaii, HI; Index of Fungi, IF; Indiana, IN; Louisiana, LA; Monoblepharidales, [M];,
Massachusetts, MA; Maryland, MD; Michigan, MI; mountain, Mt.; Neocallimastigales, [N];
North Carolina, NC; New Zealand, N.Z.; nomen nudum signifies that a taxon has no diagnosis
[S.N. Dasgupta (1982) published a "Discourse on aquatic phycomycetes of India." In appendix VI
of this paper he presented a list of aquatic fungi from studies at Lucknow. Among these are
species listed as new for which I have been unable to find descriptions. These are included herein
with the designation "? nomen nudum."]; Spizellomycetales, [S]; South America, S.A.; southeast,
SE; United Kingdom, U.K.; United States of America, USA; University, U.; Uttar Pradesh, U.P.;
Virginia, VA; Zarys Hydromikologii (Batko 1975), ZH; = indicates an opinion that taxa are
taxonomic synonyms; = indicates nomenclatural synonymy.

ACKNOWLEDGMENTS: I thank the staff of the Interlibrary Loan Department, University of
Maine Library, especially Libby Soifer and Barbara Jones, for their cooperation in obtaining copies
of journal articles; Donna Pond, Department of Plant Biology and Pathology, University of Maine
for help with printing; and Donald J.S. Barr, Centre for Land and Biological Resources Research,
Agriculture Canada for constructive suggestions and for supplying an early copy of a paper in
press. This work was encouraged and funded by Jerry R. Longcore and aided by a donation of
reprints from R.E. Reichle. I thank D.J.S. Barr and Amy Y. Rossman for helpful reviews of the
manuscript.

REFERENCES

Barr, D.J.S. 1990. Phylum Chytridiomycota. Pp. 454-466. In: Handbook of Protoctista, Eds. L.
Margulis, J.O. Corliss, M. Melkonian, and D.J. Chapman. Jones & Bartlett Publishers, Boston,
Massachusetts.

Batko, A. 1975. Zarys Hydromikologii. Paniistwowe Wydawnictwo Naukowe, Warsaw, Poland.

Dasgupta, S.N. 1982. Discourse on aquatic phycomycetes of India. Indian Phytopathol. 35: 193-
216.

Index of fungi. 1960-1995 (Vols. 3-6). C.A.B. International, Wallingford, United Kingdom.

Karling, J.S. 1977. Chytridiomycetarum Iconographia. Lubrecht & Cramer, Monticello, New
York.

Sparrow, F.K. 1960. Aquatic Phycomycetes, 2nd rev. ed. University of Michigan Press, Ann
Arbor, Michigan.

Sparrow, F.K. 1973. Chytridiomycetes, Hyphochytridiomycetes. Pp. 85-110. In: The Fungi. IVB,
Eds. G.C. Ainsworth, F.K. Sparrow, and A.S. Sussman. Academic Press, New York.

151

ALLOCHYTRIDIUM Salkin [C]; Amer. J. Bot., 57: 649-658. 1970
= Karlingia; Karling,; Nova Hedwigia, 28: 209-229. 1976; but Allochytridium
recognized by Barr & Désaulniers; see Mycologia, 78: 439-448. 1986
Allochytridium expandens Salkin, Amer. J. Bot., 57: 649-658, figs. 2-49. 1970
= Karlingia expandens (Salkin) Karling; Nova Hedwigia, 28: 209-229. 1976
Saprophytic; onion skin; roadside puddle, Solano Co., CA
Allochytridium luteum D.J.S. Barr & Désaulniers; Mycologia, 79: 193-199, figs. 1-26. 1987
Saprophytic; boiled grass; beach sand close to water level, Ontario, Canada
Allomyces; Karling (Mycopath. Mycol. Appl., 49: 169-122. 1973) established subgenera.
For review of genus see L. Olson; Opera Bot. 73: 1-96. 1984
Allomyces attomyces in S.N. Dasgupta; Indian Phytopathol. 35: 215. 1982 ? nomen nudum
Allomyces catenoides Sparrow; Mycologia, 56: 460-461. 1964 (Latin description)
For figs. see Sparrow & Morrison; Pap. Michigan Acad. Sci., 46: 175-181, 2 Pls. 1961
Saprophytic; hemp seed; aquarium with tropical fish
Allomyces reticulatus Emerson & Robertson; Amer. J. Bot., 61: 303-317, figs. 14-21. 1974
Contains key to genera of Blastocladiaceae
Saprophytic; house fly bait; cracked dry soil from drainage ditch; CA, USA
Amphicypellus Ingold 1944 = Chytriomyces Karling 1945; Dogma, I.J., Jr., Philipp. J. Biol. 5:
121-142 (p. 136) 1976. Dogma lists Chytriomyces as nomen conservandum, however,
Chytriomyces is not listed as a conserved genus in the Tokyo Code.
Amphicypellus elegans Ingold 1944 = Chytriomyces elegans (Ingold) I.J. Dogma; Philipp. J.
Biol. 5: 121-142 (p. 136). 1976
ANAEROMYCES Breton et al. [N]; FEMS Microbiol. Lett., 70: 177-182. 1990
= Ruminomyces Y.W. Ho 1990; described later, see: Mycotaxon, 47: 283-284. 1993
Anaeromyces elegans (Y.W. Ho) Y.W. Ho; Mycotaxon, 47: 283-284. 1993
= Ruminomyces elegans Y.W. Ho 1990
Anaeromyces mucronatus Breton et al.; FEMS Microbiol. Lett. 70: 177-182, fig. 1 a-f. 1990
Lectotype: see Y.W. Ho & D.J.S. Barr; Mycologia 87: 656-678. 1995
Saprophytic; hay; rumen of holstein cows; France
ARNAUDOVIA Valkanov [C]; Arch. Protistenk., 106: 553-564. 1963 (no Latin diagnosis)
= Polyphagus; Karling; CI, p. 174. 1977
Arnaudovia hyponeustonica Valkanov (no Latin diagnosis); Arch. Protistenk., 106: 553-564,
Pls. 40-43. 1963; = Polyphagus hyponeustonica (Valkanov) Karling 1977
Parasitic; Phacotus, Trachelomonas, Strombomonas spp.; ditch; Longoza, Bulgaria
Asterophlyctis irregularis Karling; Sydowia, 20: 96-108, figs. 1-16. 1967
= Septosperma irregulare (Karling) I.J. Dogma 1974
Saprophytic; Chitin bait; water containing Characeae; Lake Ohau, N.Z.
Asterophlyctis sarcoptoides H.E. Petersen 1903 = Diplophlyctis sarcoptoides (H.E. Petersen) IJ.
Dogma; Nova Hedwigia, 25: 121-141. 1974
Blastocladia arborata S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 1-3.
1988. Saprophytic; styles of submerged flowers of Caryopteris; tanks; Lucknow U., India
Blastocladia caduca S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 36-38.
1988. Saprophytic; styles of submerged flowers of Cretavea; tanks; Lucknow U., India
Blastocladia coronata S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 39-43.
1988. Saprophytic; fruit of Eugenia jambolana bait; tanks; Lucknow U., India
Blastocladia cristata in S.N. Dasgupta; Indian Phytopathol. 35: 215. 1982; ? nomen nudum
Blastocladia didyma S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, fig. 29. 1988
Saprophytic; fruit of Physalis peruviana bait; tanks; Lucknow U., India
Blastocladia elegans S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 21-22.
1988. Saprophytic; mango fruit bait; tanks; Lucknow U., Lucknow
Blastocladia excelsa S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 33-35.
1988. Saprophytic; ripe guava fruit bait; tanks; Lucknow U., India

152

Blastocladia filamentosa S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 27,
28. 1988. Saprophytic; Eugenia jambolana fruit bait; tanks; Lucknow U., India
Blastocladia fruticosa S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 4, 5.
1988. Saprophytic; Eugenia jambolana fruit bait; tanks; Lucknow U., India
Blastocladia fusiformis S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 30-32.
1988. Saprophytic; stamens and pistils of Crataeva religiosa; tanks; Lucknow U., India
Blastocladia globosa var. minutissima C. Manoharachary et al.; Bibl. Mycol., 91: 457-462, fig.
1 D. 1983. Saprophytic; unripe mango fruit bait; temple pond; Vikarabad, A.P., India
Blastocladia heterosporangia S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs.
44-45. 1988. Saprophytic; submerged grape as bait; River Gomti; Lucknow, India
Blastocladia mammilata S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs.
15-17. 1988. Saprophytic; submerged Pyrus fruit bait; tanks; Lucknow U., India
Blastocladia picaria in S.N. Dasgupta; Indian Phytopathol. 35: 215. 1982 ? nomen nudum
Blastocladia pileota S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 9-11.
1988. Saprophytic; submerged Pyrus fruit bait; tanks; garden, Lucknow U., India
Blastocladia pusilla S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 6-8. 1988
Saprophytic; mango fruit bait; tanks; Lucknow U., India
Blastocladia sessilis S.N. Dasgupta & R. John; Indian Phytopathol., 41: 521-547, figs. 23-24.
1988. Saprophytic; jamun and guava bait; tanks; Lucknow U., India
Blastocladia spiciformis S.N. Dasgupta & R. John (IF 5:946); Indian Phytopathol., 41: 521-547,
figs. 12-14. 1988. Saprophytic; guava bait; tanks; Lucknow U., India
Blastocladiella, Karling established subgenera Blastocladiella, Eucladiella & Cystocladiella;
Mycopath. Mycol. App. 49: 169-172. 1973
Blastocladiella anabaenae Canter & Willoughby; Journ. Roy. Micr. Soc., Ser. 3, 83: 365-372,
Pls. 159-162. 1964. Parasitic; Anabaena flos-aquae & A. circinalis; Lake Windermere
plankton; E.L.D., U.K.
Blastocladiella britannica Horenstein and Cantino; Trans. Brit. Mycol. Soc., 44: 185-198, Pl. 15.
1961. For figs. see Willoughby; Trans. Brit. Mycol. Soc., 42: 287-291. 1959
Saprophytic; grass, snake skin, hemp seed; periodically inundated soil; E.L.D., U.K.
Blastocladiella colombiensis Karling; Nova Hedwigia, 40: 329-340, Pls. II & Ill, figs. 4-34,
1984. Saprophytic; snake skin, insects, rarely hempseed; dry soil; La Guajira, Colombia
Blastocladiella nova-zeylandiae Karling; Sydowia, 20: 144-150, figs. 4-29. 1967
Saprophytic; bleached corn leaves and snake skin; soil; N.Z.
Blastocladiopsis elegans Robertson; Canad. J. Bot., 54: 611-615, figs. 1-7. 1976; = Ramocladia
reticulata Sorgel 1952 (nomen nudum). Saprophytic; fly; soil; Virginia Beach, VA, USA
Blyttiomyces aureus Booth; Syesis, 2: 141-161, figs. 15-22, 58-61. 1969
= B. laevis 1952; T.W. Johnson; Norw. J. Bot. 24: 83-88. 1977
Saprophytic; pine pollen; occasionally saline soil; Saturna Island, B.C., Canada
Blyttiomyces conicus T.W. Johnson; Norw. J. Bot., 24: 83-88, figs. 1-24. 1977
Saprophytic; pine pollen; water & Sphagnum; stream, Lysekloster, Norway
Blyttiomyces exuviae in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982 ? nomen nudum
Blyttiomyces gregarum in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982 ? nomen nudum
Blyttiomyces harderi Sparrow & I.J. Dogma; Arch. Mikrobiol., 89: 177-204, fig. 3, A-I. 1973
Parasitic; Rhizophlyctis rosea; soil; Santo Domingo, Dominican Republic
Blyttiomyces laevis Sparrow 1952 = Blyttiomyces aureus Booth 1969; T.W. Johnson; Norw. J.
Bot., 24: 83-88. 1977
Blyttiomyces lenis S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 42. 1988
Parasitic; zygospore of Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India
Blyttiomyces rhizophlyctidis 1.J. Dogma; Mycologia, 61: 1149-1158, figs. 1-21. 1969
Parasitic; Rhizophlyctis rosea; soil sample; Petersburg, VA, USA
Blyttiomyces spinosus S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, figs. 38-41.
1988. Saprophytic ?; Spirogyra; aquatic habitat; Lucknow, India

153

Blyttiomyces vaucheriae (Rieth) I.J. Dogma; Mycologia, 61: 1149-1158. 1969
= Phlyctochytrium vaucheriae Rieth 1956

CAECOMYCES Gold [N] in Gold et al.; BioSystems, 21: 403-415. 1988

Caecomyces communis Gold et al.; BioSystems, 21: 403-415. 1988
= Sphaeromonas communis sensu Orpin; J. Gen. Microbiol., 94: 270-280. 1976

Caecomyces equi Gold; BioSystems, 21: 403-415. figs. 1-20. 1988; ? = C. communis 1988; see
Y.W. Ho & D.J.S. Barr; Mycologia 87: 656-678. 1995
Saprophytic; plant material; caecum of horse

Callimastix cyclopis Weissenberg belongs in the Blastocladiales; see Vavra & Joyon
Protistologica, 2: 5-15 and Pls. I-VI. 1966; also Manier & Lubés, Protistologica, 4: 493-501.
1978 (not seen)

Callimastix frontalis Braune (described as protozoan; no Latin diagnosis); Arch. Protistenk., 32:
111-170, figs. 13-15. 1914 = Neocallimastix frontalis Vavra & Joyon ex Heath 1983

CANTERIA Karling [C]; Arch. Mikrobiol., 76: 126-131. 1971
based on Phlyctidium apophysatum Canter 1947

Canteria apophysata (Canter) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Phlyctidium apophysatum Canter 1947

Catenaria auxiliaris (Kiihn) Tribe; Trans. Brit. Mycol. Soc., 69: 367-376. 1977
Parasitic; Heterodera (nematodes); soil; England

Catenaria indica Singh & Pavgi; Mycologia, 62: 587-590, figs. 1-6. 1970
Saprophytic; keratin bait; soil; Varanasi, U.P., India

Catenaria ramosa W. Martin; Mycologia, 70: 461-467, figs. 3-6. 1978
Parasitic; midge eggs (Dicrotendipes modestus); streams; VA, USA

Catenaria spinosa W. Martin; Mycologia, 67: 264-272, figs. 1-15. 1975
Parasitic; chironomid eggs; small stream; VA, USA

Catenaria uncinata W. Martin; Mycologia, 70: 461-467, figs. 1,2,7,8. 1978
Parasitic; midge eggs (Glyptotendipes lobiferus); lakes and ponds; VA, USA

Catenaria vermicola Birchfield; Mycopathol. Mycol. Appl. 13: 331-338, figs. 1-3. 1960
Parasitic; plant parasitic nematodes (Xiphinema chambersi); soil; Shreveport, LA, USA

Catenaria verrucosa Karling; Sydowia, 20: 144-150, figs. 1-2. 1967
Saprophytic; snake skin bait; soil sample; N.Z.

Catenochytridium hemicysti Knox; in D.J.S. Barr et al.; Mycologia, 79: 587-594, figs. 1-25.
1987. = Chytridium hemicysta Knox; PhD dissertation, 1970; VPI, VA; not validly published
Saprophytic; cellophane, algae, pollen; Riopel's Pond; Mt. Lake Biological Station, VA, USA

Catenochytridium marinum (Kobayasi & Ookubo) Karling; CI, p. 166. 1977
= C. carolineanum f. marinum Kobayasi & Ookubo 1953

Catenochytridium oahuense Sparrow (IF 3:323); Mycopathol. Mycol. Appl., 25: 119-143, figs.
67-70, 1965. Saprophytic; cellophane bait; soil; Oahu, HI, USA

CATENOPHLYCTIS Karling [B]; Amer. J. Bot., 52: 133-138. 1965

Catenophlyctis peltata (Sparrow) Karling; Nova Hedwigia, 15: 191-201. 1968 = Phlyctorhiza
peltata Sparrow 1950

Catenophlyctis variabilis (Karling) Karling; Amer. J. Bot., 52: 133-138. 1965 = Phlyctorhiza
variabilis Karling 1947; = Entophlyctis variabilis (Karling) Cejp 1959; Omagiu lui Traian
Savulescu, 129-138. 1959

Catenophlyctis variabilis var. olduvaiensis Karling; Nova Hedwigia, 27: 747-771, figs. 48-63.
1976. Saprophytic; snake skin and corn leaves; soil; Olduvai Gorge, Tanzania

CAULOCHYTRIACEAE Subramanium; Curr. Sci., 43: 722-723. 1974. Emended by L. Olive;
Amer. J. Bot., 67: 568-574. 1986. Placed in Spizellomycetales by D.J.S. Barr; HP, p. 463. 1990

CAULOCHYTRIUM Voos & Olive [S]; Mycologia, 60: 730-733. 1968

Caulochytrium gloeosporii Voos & Olive; Mycologia, 60: 730-733, figs. 1-5. 1968; also see
Voos; Amer. J. Bot., 56: 898-909. 1969 .

Parasitic; Gloeosporium sp. on dead leguminous pods; near Miami, FL, USA

154

Caulochytrium protostelioides Olive; Amer. J. Bot., 67: 568-574, figs. 1-24. 1980
Parasitic; Cladosporium sp. on dead flowers; St. Vincent, British West Indies

Chytridiales emended by D.J.S. Barr; Canad. J. Bot., 58: 2380-2394. 1980

Chytridium Braun (in part) = Diplochytridium Karling 1971. Karling emends genus (Arch.
Mikrobiol., 76: 126-131. 1971). F.K. Sparrow (Taxon, 22: 583-586. 1973) presents figs. and
documents location of type for C. olla Braun.

Chytridium adpressum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 55. 1988
Saprophytic ?; vegetative cell of Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India

Chytridium aggregatum Karling 1938 = Diplochytridium aggregatum (Karling) Karling; Arch.
Mikrobiol., 76: 126-131. 1971

Chytridium apophysatum in S.N. Dasgupta; Indian Phytopathol, 35: 215. 1982; ? nomen nudum

Chytridium brevipes Braun 1856 = Diplochytridium brevipes (Braun) Karling; Arch.
Mikrobiol., 76: 126-131. 1971

Chytridium cejpii Fott 1950 = Diplochytridium cejpii (Fott) Karling; Arch. Mikrobiol., 76:
126-131. 1971

Chytridium chlorobotryis Fott (IF 3:38) 1952 = Diplochytridium chlorobotryis (Fott) Karling;
Arch. Mikrobiol., 76: 126-131. 1971

Chytridium citriforme Sparrow 1952 = Diplochytridium citriforme (Sparrow) Karling; Arch.
Mikrobiol., 76: 126-131. 1971

Chytridium closterii S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 66. 1988
Saprophytic ?; Closterium; tank; Lucknow U., India

Chytridium codicola Zeller 1918 = Diplochytridium codicola (Zeller) Karling; Arch.
Mikrobiol., 76: 126-131. 1971

Chytridium coleochaetes Nowakowski 1876 not Rhizophydium coleochaetes (Nowak.) Fischer
Canter; Trans. Brit. Mycol. Soc., 43: 14-18. 1960

Chytridium confervae (Wille) Minden = Chytriomyces confervae (Wille) A. Batko; ZH, p. 210.
1975

Chytridium corniculatum Kobayasi & Konno; Bull. Natl. Sci. Mus., 14: 1-18, fig. 6. 1971
Saprophytic; pine pollen; sand in fresh water; Angmagssalik, Greenland

Chytridium cresentum in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum

Chytridium deltanum Masters; Canad. J. Bot., 49: 471-481, figs. 2-4. 1971
Parasitic; Oocystis spp.; phytoplankton; Lake Manitoba, Canada

Chytridium fusiforme S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 57. 1988
Saprophytic; vegetative cell of Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India

Chytridium gibbosum Scherffel 1902 = Diplochytridium gibbosum (Scherffel) Karling; Arch.
Mikrobiol., 76: 126-131. 1971; Johnson (Nova Hedwigia, 23: 187-199. 1972 ) placed into
synonymy with Chytridium schenkii (Dangeard) Scherffel 1886.

Chytridium horariumforme S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 51.
1988. Parasitic; oospore of Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India

Chytridium hyperparasiticum Kobayasi & Konno; Bull. Natl. Sci. Mus., 14: 1-18, fig. 2 B. 1971
Parasitic; sporangium of Rhizophlyctis rosea; soil in water; Angmagssalik, Greenland

Chytridium inflatum Sparrow 1933 = Diplochytridium inflatum (Sparrow) Karling; Arch.
Mikrobiol., 76: 126-131. 1971

Chytridium isthmiophilum Canter; Trans. Brit. Mycol. Soc., 43: 660-664, fig. 1. 1960
= Diplochytridium isthmiophilum (Canter) Karling; Arch. Mikrobiol., 76: 126-131. 1971
Parasitic; Staurodesmus curvatus (desmid); plankton, lakes; E.L.D., U.K.

Chytridium kolianum Domjan 1936 = Diplochytridium kolianum (Domjan) Karling; Arch.
Mikrobiol., 76: 126-131. 1971

Chytridium lagenaria Schenk 1858 (in part) = Diplochytridium lagenarium (Schenk) Karling;
Arch. Mikrobiol., 76: 126-131. 1971

Chytridium latipodium in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum

jh

Chytridium mallomonadis Fott; Preslia, 29: 278-319, fig. 11: 1-4. 1957
= Diplochytridium mallomonadis (Fott) Karling; Arch. Mikrobiol., 76: 126-131. 1971
Parasitic; Mallomonadis tonsuratae resting spores; pond; Prague, central Bohemia

Chytridium marylandicum Paterson; Mycologia, 54: 694-703, figs. 1-7, 17 & 18. 1962
Saprophytic; matrix of Botryococcus; Blue Pond; Muirkirk, MD, USA

Chytridium mucronatum Sparrow & M. Barr 1955; = Diplochytridium mucronatum (Sparrow
& M. Barr) Karling; Arch. Mikrobiol., 76: 126-131. 1971

Chytridium neopapillatum Kobayasi & Konno; Bull. Natl. Sci. Mus., 14: 1-18, fig. 7 A. 1971
Saprophytic; pine pollen; sand in water; Angmagssalik, Greenland

Chytridium oedogonii Couch 1938 = Diplochytridium oedogonii (Couch) Karling; Arch.
Mikrobiol., 76: 126-131. 1971

Chytridium ottariense Roane (IF 4:241); Mycologia, 65: 531-538, figs. 3-14. 1973
Saprophytic; Volvox, chitin, snake skin; impoundment; Pulaski County, VA, USA

Chytridium parasiticum Willoughby 1956; = Chytriomyces willoughbyi (Willoughby) Karling;
Mycopath. Mycol. Appl., 36: 165-178. 1968

Chytridium pilosum Kobayasi & Konno; J. Jap. Bot., 45: 325-337, fig. 2 A-K. 1970
Saprophytic; cellophane bait; soil from evergreen forest; Tsushima Island, Japan

Chytridium proliferum Karling; Sydowia, 20: 119-128, figs. 16-28. 1967
Saprophytic; pollen, corn leaves, snake skin; acidic soil; N.Z.

Chytridium reniforme S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 54. 1988
Saprophytic ?; Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India

Chytridium schenkii (Dangeard) Scherffel 1886 emended by T.W. Johnson; Nova Hedwigia, 23:
187-199. 1972 = Chytridium gibbosum Scherffel 1902
= Diplochytridium schenkii (Dangeard) Karling; Arch. Mikrobiol., 76: 126-131. 1971

Chytridium schenkii var. dumontii 1.J. Dogma; Philipp. Agric., 57: 49-54, figs. 1-21. 1973
Saprophytic; cellophane, lens paper, onion skin; moist soil; Columbia, S.A.

Chytridium scherffelii (Scherffel) Sparrow 1936
= Diplochytridium scherffelii (Sparrow) Karling; Arch. Mikrobiol., 76: 126-131. 1971

Chytridium sexuale Koch 1951 = Diplochytridium sexuale (Koch) Karling; Arch. Mikrobiol.,
76: 126-131. 1971

Chytridium sparrowii S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 56. 1988
Saprophytic ?; Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India

Chytridium stellatum (Petersen) Koch 1951 = Diplochytridium stellatum (Petersen) Karling;
Arch. Mikrobiol., 76: 126-131. 1971

Chytridium telmatoskenae Fott; Preslia, 29: 278-319, fig. 11: 5-10. 1957
Parasitic; Telmatoskene mucosa; pond; Volary, central Bohemia

Chytridium turbinatum Kobayasi & Ookubo 1954 = Diplochytridium turbinatum (Kobayasi
& Ookubo) Karling; Arch. Mikrobiol., 76: 126-131. 1971

Chytriomyces Karling 1945 includes Amphicypellus Ingold 1944; I.J. Dogma: Philipp. J. Biol. 5:
121-142 (p. 136). 1976. Dogma lists Chytriomyces as "nomen conservandum"; however
Chytriomyces is not on the conserved list in the Tokyo Code.

Chytriomyces angularis Longcore; Mycologia, 84: 442-451, figs. 1-28. 1992
Saprophytic; pollen, snake skin; lakes, Sphagnum; Maine, USA

Chytriomyces annulatus 1.J. Dogma; Nova Hedwigia, 18: 349-365, figs. 1-18. 1969

_ Saprophytic; pollen, snake skin; leaf litter and forest soils; USA

Chytriomyces confervae (Wille) A. Batko; ZH, p. 210. 1975 = Chytridium confervae (Wille)
Minden 1915

Chytriomyces cosmarii Karling; Sydowia, 20: 119-128, figs. 1-7. 1967
Parasitic; Cosmarium spp.; soil, hillside sheep paddock; Taita, N.Z.

Chytriomyces elegans (Ingold) I.J. Dogma; Philipp. J. Biol. 5: 121-142 (p. 136). 1976
= Amphicypellus elegans Ingold 1944

Chytriomyces gilgaiensis Willoughby; Arch. Mikrobiol., 52: 101-131, fig. 6 a-l. 1965
Parasitic; Nowakowskiella crassa; soil, gilgai depression; Victoria, Australia

156

Chytriomyces heliozoicola Canter; Trans. Brit. Mycol. Soc., 49: 633-638, fig. 1. 1966
Parasitic; Raphidiocystis lemani; plankton of lakes; E.L.D., U.K.

Chytriomyces hyalinus Karling emended by Bostick; J. Elisha Mitchell Sci. Soc. 84: 94-99. 1968

Chytriomyces hyalinus var. granulatus Karling; Sydowia, 20: 119-128, figs. 8-15. 1967
Saprophytic; corn leaves, chitin, and snake skin; soil; widespread in N.Z.

Chytriomyces laevis Karling; Nova Hedwigia, 44: 137-139, figs. 1-18. 1987
Parasitic; Pythium; soil, stream bank; Panama

Chytriomyces macro-operculatus Karling; Nova Hedwigia, 34: 645-668, figs. 24-38, 1981
Saprophytic; cellophane & corn cotyledons; sandy organic soil; Isla Orquidia, Venezuela

Chytriomyces macro-operculatus var. hirsutus Karling; Nova Hedwigia, 34: 645-668. figs.
39-45. 1981. Saprophytic; cellophane; sandy organic soil; Isla Orquidia, Venezuela

Chytriomyces mammilifer Persiel; Arch. Mikrobiol., 36: 283-305, figs. 9, 10. 1960
Saprophytic; pine pollen; soil from altitude of 4 K; Ecuador

Chytriomyces mortierellae Persiel; Arch. Mikrobiol., 36: 283-305, figs. 6, 7. 1960
Parasitic; Mortierella; soil sample; summit of Piz Buin, Austria

Chytriomyces multi-operculatus Sparrow & 1.J. Dogma; Arch. Mikrobiol., 89: 177-204, fig. 5.
1973. Saprophytic; pine pollen; shallow, black soil; Dominican Republic

Chytriomyces nagatoroensis Konno; Sci. Rep. Tokyo Kyoiku Daigaku Sect. B, 14: 227-292, PI.
4, fig. X. 1972. Parasitic-weak; Spirogyra sp.; water?; Nagatoro (Saitama Prefecture), Japan

Chytriomyces poculatus Willoughby & Townley; Trans. Brit. Mycol. Soc., 44: 177-184, fig. 3,
Pl. 14.1961. Saprophytic; snake skin; soil; E.L.D., U.K.

Chytriomyces reticulatus Persiel; Arch. Microbiol., 36: 283-305, fig. 8. 1960
Parasitic; Pythium proliferum, soil from flower pot; Gottingen, Germany

Chytriomyces reticulosporus 1.J. Dogma; Philipp. J. Biol., 12: 385-408, figs. 10-21. 1983
Parasitic; Phlyctochytrium punctatum, cultivated soils; Luzon, Philippines

Chytriomyces rhizidiomycetis 1.J. Dogma (IF 5:282); Philipp. J. Biol., 12: 385-408, figs. 1-9.
1983. Parasitic; Rhizidiomyces bivellatus; cultivated soils; Philippines & Japan

Chytriomyces rotoruaensis Karling; Arch. Microbiol., 70: 266-287, figs. 1 P-Y, 2 A-S. 1970
Saprophytic; purified shrimp chitin; mud, cooled thermal pool; Rotorua, N.Z.

Chytriomyces suburceolatus (Willoughby) Willoughby; Nova Hedwigia, 7: 133-150. 1964
= Chytridium suburceolatum Willoughby 1956

Chytriomyces vallesiacus Persiel; Arch. Mikrobiol., 36: 283-305, fig. 11. 1960
Saprophytic; pine pollen; soil; "Felsband am Rimpfischhorn im Wallis," Austria

Chytriomyces verrucosus Karling; Bull. Torrey Bot. Club, 87: 326-336, figs. 19A-K. 1960
Parasitic; Karlingia rosea; soil; Lafayette, IN, USA

Chytriomyces willoughbyi (Willoughby) Karling; Mycopathol. Mycol. Appl., 36: 165-178. 1968
= Chytridium parasiticum Willoughby 1956

Cladochytrium aureum Karling 1949 = C. replicatum Karling 1931; Karling; Sydowia, 20: 129-
136. 1967

Cladochytrium granulatum (Karling) Sparrow 1960 = Nowakowskiella granulata Karling;
Sydowia, Beiheft VI, p. 66. 1966

Cladochytrium indicum Singh & Pavgi; Hydrobiologia, 37: 565-568, figs. 1-7. 1971
Saprophytic; snake skin bait; soil; Deoria, U.P., India

Cladochytrium novoguineense Kobayasi & Konno; Bull. Natl. Sci. Mus., 14: 373-386, fig. 3
C-I. 1971. Saprophytic; cellophane and onion skin; soil; Papua, New Guinea & Solomon
Islands

Cladochytrium replicatum Karling 1931 = C. aureum Karling 1949; Karling; Sydowia, 20:
129-136. 1967

Cladochytrium salsuginosum A. Batko & S.K.M. Hassan; Acta Mycol., 22: 189-192, figs. 1-9.
1986 (1988); Saprophytic; onion skin bait; brackish water; near Frombork, northern Poland

Coelomomycetaceae; new spp.not included, see “The genus Coelomomyces’’, Eds. J.N. Couch
& C.E. Bland, Academic Press, Inc., New York. 1985

Coelomycidium, placed in Blastocladiales; D.J.S. Barr HP, p. 463. 1990

157

Cylindrochytrium endobioticum Willoughby; Nova Hedwigia, 7: 133-150, Pls. 17, 18 a-d, 23.
1964. Saprophytic; cellophane and grass leaf baits; soil; Bogle Crag Wood, Lancashire, U.K.
Cystocladiella Karling (subgenus of Blastocladiella); Mycopath. Mycol. Appl. 49: 169-172. 1973
Dangeardia appendiculata (Zopf) Batko; Acta Mycol., 6: 407-435. 1970 (contains key to
Dangeardia) = Scherffeliomyces appendiculatus (Zopf) Sparrow 1936
Dangeardia echinulata A. Batko; Acta Mycol., 6: 407-435, figs. 6-20. 1970
Parasitic; Vitreochlamys (Sphaerellopsis) aulata; pool near Vistula River; Warsaw, Poland
Dangeardia molesta (Canter) A. Batko; ZH, p. 179. 1975 = Entophlyctis molesta Canter 1965
Dangeardia ovata Paterson; Mycologia, 50: 453-468, figs. 1-6. 1958
Parasitic; Sphaerocystis schroeteri;, plankton, fakes; Washtenaw Co., MI, USA
Dangeardia sporapiculata Geitler; Sydowia, 16: 324-330, figs. 1-2. 1963
Parasitic; Chlamydomonas palmaloid stage; lake; near Lunz, Austria
Dangeardia sporapiculata var. minor Geitler (? no type); Oesterr. Bot. Z., 112: 603-609. 1965
Parasitic; Heleochloridis pallidae; soil; Schladming, Austria
DANGEARDIANA Valkanov ex A. Batko [C]; Acta Mycol., 6: 407-435, 1970; includes key to
related forms. = Dangeardiana Valkanov (no Latin diagnosis); Arch. Mikrobiol., 48: 239-246.
1964. Karling doubted need to separate from Dangeardia (CI, p. 100. 1977).
Dangeardiana apiculata (Braun) A. Batko; Acta Mycol., 6: 407-435. 1970
= Entophlyctis apiculata (Braun) Fischer (1892); also see Karling: CI p. 100. 1977
Dangeardiana eudorinae Valkanov ex A. Batko; Acta Mycol., 6: 407-435. 1970
= Dangeardiana eudorinae Valkanov (no Latin diagnosis); Arch. Mikrobiol., 48: 239-246, figs.
1-12. 1964. Parasitic; oospores of Eudorina elegans; water; Sofia, Bulgaria
Dangeardiana leptorrhiza (Johns) A. Batko; Acta Mycol., 6: 407-435. 1970
= Scherffeliomyces leptorrhizus Johns 1956
Dangeardiana sporapiculata (Geitler) A. Batko; Acta Mycol., 6: 407-435. 1970
= Dangeardia sporapiculata Geitler 1963
DICTYOMORPHA Mullins [C]; Amer. J. Bot., 48: 377-387. 1961 = Pringsheimiella Couch
1939 (invalid homonym)
Dictyomorpha dioica Couch ex Mullins; Amer. J. Bot., 48: 377-387, figs. 1-37. 1961
= Pringsheimiella dioica Couch 1939. Parasitic; Oomycetes; Mirror Lake; Highlands, NC, USA
Dictyomorpha dioica var. pythiensis N. Sarkar & R. Dayal; Proc. Natl. Acad. Sci. India, Sect. B,
58: 403-406, figs. 1-10. 1988 .
Parasitic; Pythium aphanidermatum; Kurukshetra Pond; Varanasi, U.P., India
DIPLOCHYTRIDIUM Karling [C]; Arch. Mikrobiol., 76: 126-131. 1971
Segregate of Chytridium Braun
Diplochytridium aggregatum (Karling) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium aggregatum Karling 1938
Diplochytridium brevipes (Braun) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium brevipes Braun 1856
Diplochytridium cejpii (Fott) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium cejpii Fott 1950
Diplochytridium chlorobotryis (Fott) Karling (IF 3:62); Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium chlorobotytris Fott 1952
Diplochytridium citriforme (Sparrow) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium citriforme Sparrow 1952
Diplochytridium codicola (Zeller) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium codicola Zeller 1918
Diplochytridium gibbosum (Scherffel) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium gibbosum Scherffel 1926
Diplochytridium inflatum (Sparrow) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium inflatum Sparrow 1933
Diplochytridium isthmiophilum (Canter) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium isthmiophilum Canter 1960

158

Diplochytridium kolianum (Domjan) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium kolianum Domjan 1936

Diplochytridium lagenarium var. japonense (Kobayasi & Ookubo) Karling; Arch. Mikrobiol.,
76: 126-131. 1971; = Chytridium lagenaria var. japonense Kobayasi & Ookubo 1953

Diplochytridium lagenarium (Schenk) Karling (IF 4:62); Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium lagenaria Schenk 1858 (pro parte)

Diplochytridium mallomonadis (Fott) Karling,; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium mallomonadis Fott 1957

Diplochytridium mucronatum (Sparrow & M. Barr) Karling; Arch. Mikrobiol., 76: 126-131.
1971; = Chytridium mucronatum Sparrow and M. Barr 1955

Diplochytridium oedogonii (Couch) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium oedogonii Couch 1938

Diplochytridium schenkii (Scherffel) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium schenkii Scherffel 1926

Diplochytridium scherffelii (Sparrow) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium scherffelii Sparrow 1936

Diplochytridium sexuale (Koch) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium sexuale Koch 1951

Diplochytridium stellatum (Petersen) Karling; Arch. Mikrobiol., 76: 126-131. 1971
= Chytridium stellatum (Petersen) Koch 1951

Diplochytridium? turbinatum (Kobayasi & Ookubo) Karling; Arch. Mikrobiol., 76: 126-131.
1971; = Chytridium turbinatum Kobayasi & Ookubo 1954

Diplophlyctis asteroidea 1.J. Dogma & key to chitinophilic Diplophlyctis spp.; Trans. Brit.
Mycol. Soc., 67: 255-264, figs.1-3. 1976; Saprophytic; chitin; acid bogs; northern MI, USA

Diplophlyctis buttermerensis (Willoughby) Dogma (IF 3:420); Arch. Mikrobiol., 66: 203-219,
1969; = Nephrochytrium buttermerense Willoughby 1962
A. Batko retains N. buttermerense; ZH, p. 220. 1975

Diplophlyctis chitinophila Willoughby; Trans. Brit. Mycol. Soc., 45: 121-136, fig. 5. 1962
Saprophytic; chitin; submerged mud; lakes, E.L.D., U.K.

Diplophlyctis complicata (Willoughby) Dogma (IF 3:306); Nova Hedwigia, 25: 143-159. 1974
= Nephrochytrium complicatum Willoughby 1961

Diplophlyctis nephrochytrioides Karling; Sydowia, 20: 109-118, figs. 36-67. 1967
Saprophytic?; Nitella, moribund internodes; Lake Ohau; Canterbury Province, N.Z.

Diplophlyctis sarcoptoides (Petersen) I.J. Dogma; Nova Hedwigia, 25: 121-141. 1974
= Asterophlyctis sarcoptoides Petersen 1903
Dogma (/oc.cit.) designated figs. 1-38 on Pls. I-IV as neotype.

Diplophlyctis sexualis Haskins 1950 = Nephrochytrium sexuale (Haskins) A Batko; ZH,
p. 220. 1975

Diplophlyctis versiformis 1.]. Dogma; Philipp. J. Biol., 5: 121-142, figs. 1-25. 1976
Saprophytic; shrimp chitin; Bryant's Bog, Smith's Bog; MI, USA

Endochytrium cystarum 1.J. Dogma; Arch. Mikrobiol., 66: 203-219, figs. 26-39. 1969
Saprophytic; lens paper and cellophane bait; forest soil and leaf litter; MI, USA

Endochytrium multiguttulatum 1.J. Dogma; Arch. Mikrobiol., 66: 203-219, figs. 1-25. 1969
Saprophytic; lens paper, cellophane, onion; bogs; Cheboygan Co., MI, USA

Entophlyctis Fischer 1892; D.J.S. Barr (Canad. J. Bot., 58: 2380-2394. 1980.) emended and
placed in Spizellomycetales; Longcore et al. (Canad. J. Bot., 73: 1385-1390. 1995) rejected
emendment and described Powellomyces for Entophlyctis (sensu Barr, 1980);
Entophlyctis (sensu Fischer) is in Chytridiales, see Longcore; Mycologia, 87: 25-33. 1995

Entophlyctis apiculata (Braun) Fischer 1892 = Dangeardiana apiculata (Braun) A. Batko;
Acta Mycol., 6: 407-435. 1970

Entophlyctis bulligera (Zopf) Fischer (1884) = Entophlyctis confervae-glomeratae
(Cienkowski) Sparrow (1857); D.J.S. Barr: Canad. J. Bot., 49: 2215-2222. 1971

159

Entophlyctis bulligera var. brevis S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig.
44. 1988; Saprophytic ?; vegetative Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India
Entophlyctis caudiformis S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 47.
1988. Saprophytic ?; Closterium; tank; Lucknow U., India
Entophlyctis confervae-glomeratae (Cienkowski) Sparrow (1857); D.J.S. Barr (Canad. J. Bot.,
49: 2215-2222. 1971) considers the following synonyms: = E. bulligera (Zopf) Fischer;
= E. vaucheriae; (Fisch) Fischer, = E. helioformis (Dangeard) Ramsbottom;
= E. rhizina (Schenk) Minden. Powell & Koch (Canad. J. Bot., 55: 1668-1685. 1977) do not
consider E. bulligera a synonym.
Entophlyctis crenata Karling; Sydowia, 20: 109-118, figs. 1-12. 1967
Saprophytic; Vallisneria sp.; small brook; Wellington Province, N.Z.
‘ Entophlyctis filamentosa in S.N. Dasgupta; Indian Phytopathol. 35: 215. 1982; ? nomen nudum
Entophlyctis helioformis (Dangeard Ramsbottom (1886); = Entophlyctis confervae-glomeratae
(Cienkowski) Sparrow (1857); D.J.S. Barr: Canad. J. Bot., 49: 2215-2222. 1971
Entophlyctis lobata Willoughby & Townley; Trans. Brit. Mycol. Soc., 44: 177-184. figs. 1-2, PI.
13. 1961. Saprophytic; shrimp chitin bait; littoral of Derwentwater; E.L.D., U.K.
Entophlyctis luteolus Longcore; Mycologia, 87: 25-33, figs. 1-20. 1995
Saprophytic; onion skin and Utricularia; bogs and acid lake; Maine (also MI), USA
Entophlyctis mammilliformis S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 46.
1988. Saprophytic ?; oospores of Spirogyra; pond; Mohanlalganj, east of Lucknow U., India
Entophlyctis molesta Canter; J. Roy. Microscop. Soc. London, 84: 549-557, figs. 1-4. 1965
= Dangeardia molesta (Canter) A. Batko 1975
?Parasitic; Stylosphaeridium stipitatum; phytoplankton; English lakes, U.K.
Entophlyctis obscura Sparrow & I.J. Dogma; Arch. Mikrobiol., 89: 177-204, fig. 2 A-S. 1973
Saprophytic; pine pollen bait; soil; Dominican Republic
Entophlyctis reticulospora P. Cook; Trans. Brit. Mycol. Soc., 49: 545-550, Pls. 28-29. 1966
Parasitic; Closterium moniliferum et al.; swampy area; Cheboygan Co., MI, USA
Entophlyctis rhizina (Schenk) Minden (1858) = Entophlyctis confervae-glomeratae
(Cienkowski) Sparrow (1857); D.J.S. Barr; Canad. J. Bot., 49: 2215-2222. 1971
Entophlyctis sphaerioides S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 48.
1988. Saprophytic ?; reproductive body of Spirogyra; ditch; Sultanpur Road, SE of Lucknow
U., India
Entophlyctis texana Karling 1941; see Sydowia, 20: 109-118. 1967 for 1st published figs.
Entophlyctis variabilis (Karling) Cejp; Omagiu lui Traian Savulescu, 129-138. 1959
= Phlyctorhiza variabilis Karling 1947; = Catenophlyctis variabilis (Karling) Karling 1965
Entophlyctis variabilis Powell & Koch [not E. variabilis (Karling) Cejp 1959] (homonym)
= Powellomyces variabilis Powell & Koch ex Longcore et al. 1995
Saprophytic; pollen and Paspalum leaf; soils; NC, USA
Entophlyctis vaucheriae (Fisch) Fischer 1884; = Entophlyctis confervae-glomeratae
(Cienkowski) Sparrow 1857; D.J.S. Barr; Canad. J. Bot., 49: 2215-2222. 1971
Entophlyctis willoughbyi Bradley (fossil); Amer. J. Bot., 54: 577-582, figs. 1-7, 9. 1967
In fossil plant parts; Eocene Green River Formation, Wyoming, USA
Eucladiella Karling (subgenus of Blastocladiella); Mycopath. Mycol. App. 49: 169-172. 1973
GAERTNERIOMYCES D.J.S. Barr [S]; Canad. J. Bot., 58: 2380-2394. 1980
Gaertneriomyces semiglobiferus (Uebelmesser) D.J.S. Barr; Canad. J. Bot., 58: 2380-2394. 1980
= Phlyctochytrium semiglobiferum Uebelmesser 1956; = P. californicum D.J.S. Barr, 1969
Gaertneriomyces tenuis D.J.S. Barr, Canad. J. Bot., 62: 1171-1201, figs. 129-137. 1984
Saprophytic; pollen; soil; near Port William Dyke, Nova Scotia, Canada
Gonopodya terrestris S.N. Dasgupta et al, ( probably not a Gonopodya); Indian Phytopathol., 43:
218-222, Figs. 1-8. 1990. Saprophytic; leaves of Eugenia jambolana; moist soil; garden,
Lucknow U., India
Hapalopera Fott 1942; accepted by A. Batko; ZH, Pp. 169-170. 1975; rejected by Karling; CI,
Pp. 64, 65. 1977

160

Hapalopera achnanthis (Friedmann) A. Batko;, ZH, p. 170. 1975 = Rhizophydium achnanthis
Friedmann 1952

Hapalopera difficilis (Canter) A. Batko; ZH, p. 169. 1975 = Rhizophydium difficile Canter 1954

Hapalopera fragilariae (Canter) A. Batko, ZH, p. 170. 1975 = Rhizophydium fragilariae Canter
1950

Hapalopera melosirae (Friedmann) A. Batko; ZH, p. 170. 1975 = Rhizophydium melosirae
Friedmann 1952

Hapalopera piriformis Fott; A. Batko; ZH, p. 170. 1975 = Rhizophydium piriformis (Fott)
Karling; CI. p. 65. 1977

HARPOCHYTRIACEAE Wille; Petermann's Geog. Mitt. Erganzungsband, 28: 370-371. 1900
Emended by Emerson & Whisler; Arch. Mikrobiol., 61: 195-211. 1968
Placed in Chytridiales by D.J.S. Barr; HP, p.463. 1990

HARPOCHYTRIALES Emerson & Whisler; Arch. Mikrobiol., 61: 195-211. 1968
Not used by D.J.S. Barr; HP, p.463. 1990; used by Powell & Roychoudhury; Canad. J. Bot., 70:
750-761. 1992

HARPOCHYTRIUM Lagerheim; Hedwigia, 29: 142-145. 1890
Placed in Harpochytriales by Emerson & Whisler; Arch. Mikrobiol., 61: 195-211. 1968
From Harpochytriales to Chytridiales; Barr; HP, p.463. 1990

Harpochytrium adpressum Scherffel; Arch. Protistenk., 54: 510-528, Pl. 28, figs.17, 18. 1926
Saprophytic; Spirogyra jugalis; water near igloo

Harpochytrium apiculatum Pascher (paper not seen) see: Pascher; Heterokonten, Lief. 5 in
Rabenhorst's Kryptogamen-Flora von Deutschland II. Leipzig. 1938

Harpochytrium botryococci Jane; J. Linn. Soc., Bot., 53: 28-40, figs. 14-18. 1946
Saprophytic; in mucilage of Botryococcus; water; Llyn Bychan, Caernarvonshire, U.K.

Harpochytrium hedenii Wille; Petermann's Geogr. Mitt. Erganzungsband 28: 370-371. 1900
Saprophytic; Zygnema, Oedogonium, Spirogyra;

Harpochytrium hyalothecae Lagerheim; Hedwigia, 29: 142-145, figs. 1-4. 1890
On Hyalotheca dissiliens; "Lassby backar"; Upsala, Sweden

Harpochytrium intermedium Atkinson; Ann. Mycol., 1: 479-502, Pl. 10, fig. 22, a 1903
Saprophytic; Confervae; marsh; near Ithaca, New York, USA
F.W. Jane considers this species a nomen nudum; J. Linn. Soc., Bot., 53: 28-40. 1946

Harpochytrium monae Jane; J. Linn. Soc., Bot., 53: 28-40, figs. 19-22. 1946
Saprophytic; Zygnema; temporary pool; Llandegfan Common, Anglesey, U.K.

Harpochytrium natrophilum Hortobagyi (alga); Bot. K6zlem., 48: 217-223. 1960

Harpochytrium ornithocephalum Clémencon; Rev. Algol., N.S., 9: 33-37, figs. 12-20 (Pl. 5).
1969. Saprophytic; on Spirogyra; fishpond; Botanical garden, Bern, Switzerland

Harpochytrium tenuissimum Korschikoff; Arch. Protistenk., 74: 249-258, figs. 21-22. 1931
Saprophytic; On Oedogonium sp.; Latzino-Moores; Zwenigorod, Germany
Emended by Jane; J. Linn. Soc., Bot., 53: 28-40. 1946

Harpochytrium viride Scherffel (alga); Arch. Protistenk., 54: 510-528. 1926

Karlingia Johanson 1944; Sparrow 1960 placed exo-operculate spp. in Karlingiomyces.
Karlingiomyces not accepted by Karling; Sydowia, Beiheft VI., p. 58. 1966

Karlingia aurantiaca Karling; Nova Hedwigia, 27: 747-771, figs. 22-31. 1976
Saprophytic; corn leaves; soil; Blantyre, Malawi

Karlingia exo-operculata Karling; Nova Hedwigia, 28: 209-229, figs. 1-16. 1976
Saprophytic; cellophane bait; soil; Orlando, FL, USA

Karlingia expandens (Salkin) Karling; Nova Hedwigia, 28: 209-229. 1976
= Allochytridium expandens Salkin 1970; A. expandens preferred.

Karlingia granulata Karling not Karlingiomyces granulatus (Karling) Sparrow 1960
IJ. Dogma; Nova Hedwigia, 25: 91-105. 1974

Karlingia lacustris Hassan; Nova Hedwigia, 37: 435-440, figs. 1-7. 1983
Saprophytic; onion skin bait; Mikolajki Lake; northeast Poland

161

Karlingia lobata var. microspora Karling; Nova Hedwigia, 40: 329-340, Pl. 1, figs. 1-18. 1984
Saprophytic; bleached corn cotyledons; soil; LaGuajira, Colombia, S.A.
Karlingia polonica Hassan; Nova Hedwigia, 38: 727-740, figs.1-17, Pl. 1. 1983
Saprophytic; onion skin; water, Mlociny Park, Poland
Karlingia spinosa Karling not Rhizophlyctis spinosa (Karling) Sparrow 1960
I.J. Dogma; Nova Hedwigia, 25: 91-105. 1974
Karlingiomyces Sparrow 1960; rejected by Karling; Sydowia, Beiheft VI. p. 58. 1966
Accepted by I.J. Dogma; Nova Hedwigia, 24: 393-411. 1973
Karlingiomyces laevis Konno; Sci. Rep. Tokyo Kyoiku Daigaku Sect. B, 14: 227-292, Pl. 8,
E-H. 1972. Saprophytic; cellophane bait; forest soil; Susono-machi (Shizuoka Prefecture),
Japan
KOCHIOMYCES D.J.S. Barr [S]; Canad. J. Bot., 58: 2380-2394. 1980
Kochiomyces dichotomus (Umphlett) D.J.S. Barr (IF 5:51); Canad. J. Bot., 58: 2380-2394. 1980
= Phlyctochytrium dichotomum Umphlett 1967
LACUSTROMYCES Longcore [C]; Canad. J. Bot., 71: 414-425. 1993
Lacustromyces hiemalis Longcore; Canad. J. Bot., 71: 414-425, figs. 1-42. 1993
Saprophytic; chitin; lakes; Maine, USA
Macrochytrium botrydiella in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? Nom. nudum
Macrochytrium botrydioides var. minutum S.N. Dasgupta & R. John (no Latin diagnosis); Bull.
Bot. Surv. India, 30: 1-82, fig. 67. 1988.
Saprophytic; submerged pears as bait; tank; Lucknow U., India
MICROALLOMYCES Emerson & Robertson [B]; Amer. J. Bot., 61: 303-317. 1974
Microallomyces dendroideus Emerson & Robertson; Amer. J. Bot., 61: 303-317, figs. 1-13.
1974. Saprophytic; house fly bait; damp soil from dried pond; Costa Rica
Micromyces furcata Rieth; Kulturpflanze, Beiheft 3: 286-295, Pl. 2 A-H. 1962
Parasitic; Spirogyra longata; Stechlinsee region; Germany
Micromyces grandis Miller 1955 = Synchytrium grandis (Miller) Karling; "Synchytrium",
Academic Press, NY. p. 124. 1964
MICROMYCOPSIDACEAE Subramanium; Curr. Sci., 43: 722-723. 1974
Family not included by D.J.S. Barr; HP, p. 463, 1990.
Mitochytridium regale S.K.M. Hassan; Acta Mycol., 18: 155-160, figs.1-4, Pl. I. (1982) 1986
Saprophytic; snake skin bait; pond; Lazienki Royal Garden, Warsaw, Poland
Monoblepharella placed in Monoblepharidaceae by A. Batko; ZH, p. 249-250. 1975
Also placed in Monoblepharidaceae, perhaps accidentally, by D.J.S. Barr, HP, p.463. 1990
Monoblepharis micrandra Sparrow; Trans. Brit. Mycol. Soc., 48: 55-58, fig. 1 A-G. 1965
Saprophytic; twigs; shrubby swamp; Rifle River Recreation Area, MI, USA
Monophagus in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum
Monophagus blackmanii in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum
Monophagus bruhlii in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum
NEOCALLIMASTIGACEAE Heath (IF) [N]; Canad. J. Bot., 61: 295-307. 1983
Emended by Barr et al.; Canad. J. Bot., 67: 2815-2824. 1989
NEOCALLIMASTIGALES J. Li, Heath, & Packer (IF); Canad. J. Bot., 71: 393-407. 1993
Review: Li & Heath; Canad. J. Microbiol. 39: 1003-1013. 1993
Review: Trinci et al.; Mycol. Res. 98: 129-152. 1994
Taxonomy: Y.W. Ho & D.J.S. Barr; Mycologia, 87: 656-678. 1995
NEOCALLIMASTIX Vavra & Joyon ex Heath [N]; Canad. J. Bot., 61: 295-307. 1983
= Neocallimastix Vavra & Joyon (no Latin) ; Protistologica, 2: 5-15, Pl. 1-6. 1966
Emended by Orpin & Munn; Trans. Brit. Mycol. Soc., 86: 178-181, figs. 1-4. 1986
Neocallimastix frontalis (Braune) Vavra & Joyon ex Heath; Can. J. Bot., 61: 295-307. 1983
= Callimastix frontalis Braune 1913. Lectotype: see Y.W. Ho & D.J.S. Barr; Mycologia 87:
656-678. 1995. Y.W.Ho & DJ.S. Barr (loc. cit.) place the following in synonymy:
=N. variabilis Y.W. Ho & D.J.S. Barr 1993; = N. patriciarum Orpin & E.A. Munn 1986

162

Neocallimastix hurleyensis Theodorou & J. Webb; Canad. J. Bot., 69: 1220-1224, figs. 1-7.
1991. ? = Neocallimastix frontalis (Braune) Heath; D.J.S. Barr: Mycotaxon, 54: 203-214. 1995
Saprophytic; plant materials; rumen of sheep; U.K.

Neocallimastix joyonii Breton et al.; FEMS Microbiol. Lett., 58: 309-314, figs. 1-2. 1989
= Orpinomyces joyonii (Breton et al.) Li et al. 1991
Saprophytic; straw; rumen of sheep; France.

Neocallimastix patriciarum Orpin & Munn, Trans. Brit. Mycol. Soc., 86: 178-181, figs. 1-4.
1986. = Neocallimastix frontalis (Braune) Heath; D.J.S. Barr: Mycotaxon, 54: 203-214. 1995
Saprophytic; plant tissues; rumen of sheep

Neocallimastix variabilis Y.W. Ho & D.J.S. Barr; Mycotaxon, 46: 241-258, figs. 1-10. 1993
= Neocallimastix frontalis (Braune) Heath; D.J.S. Barr: Mycotaxon, 54: 203-214. 1995
Saprophytic; guinea grass (Panicum maximum), rumen of Bos indicus; peninsular Malaysia

Nephrochytrium bipes Hassan; Nova Hedwigia, 38: 727-740, figs. 18-28, Pl. 2. 1983
Saprophytic; onion skin; pond; near Warsaw, Poland

Nephrochytrium buttermerense Willoughby; Nova Hedwigia, 3: 439-444, Pls. 114-116. 1961
= Diplophlyctis buttermerense (Willoughby) I.J. Dogma 1969
Not D. butermerense (Willoughby) I.J. Dogma; A. Batko; ZH, p. 220. 1975
Saprophytic; cellophane bait; submerged mud; Buttermere, E.L.D., U.K.

Nephrochytrium complicatum Willoughby; Nova Hedwigia, 3: 439-444, Pls. 112, 113. 1961
= Diplophlyctis complicata (Willoughby) I.J. Dogma 1974;
combination also made by A. Batko; ZH, p. 186. 1975
Saprophytic; chitin bait; submerged lake mud; Ennerdale, E.L.D., U.K.

Nephrochytrium sexuale (Haskins) A. Batko; ZH, p. 220. 1975 = Diplophlyctis sexualis
Haskins 1950

Nowakowskiella crassa Karling 1949; = Nowakowskiella elegans (Nowakowski) Schroeter
1893. See T.W. Johnson; Mycologia, 69: 34-45, 1977

Nowakowskiella delica Whiffen 1943 = Nowakowskiella elegans (Nowakowski) Schroeter 1893
See T.W. Johnson; Mycologia, 69: 34-45, 1977

Nowakowskiella elegans (Nowakowski) Schroeter 1893 = N. crassa Karling 1949,
= N. delica Whiffen 1943, = N. endogena Constantineau 1901, and = N. profusa Karling 1944
See T.W. Johnson; Mycologia, 69: 34-45. 1977

Nowakowskiella granulata Karling 1944 = Cladochytrium granulatum (Karling) Sparrow 1960
Karling (Sydowia, Beiheft VI., p. 66. 1966) did not agree with placement in Cladochytrium.

Nowakowskiella keratinophila S.K.M. Hassan & A. Batko; Acta Mycol., 22: 193-196, figs. 1-7.
1986 (1988). Saprophytic; snake skin bait; brackish water; Bay Zalew Wislany, Poland

Nowakowskiella methistemichroma A. Batko & S.K.M. Hassan; & key to Nowakowskiella spp.
Sydowia, 35: 27-36, figs. 1-21, 1982
Saprophytic; onion skin bait; water and mud; Kampinos National Park, Poland

Nowakowskiella moubasherana S.K.M. Hassan (IF 5:397); Acta Mycol., 19: 77-82, figs. 1-18
& Pl. I. 1983. Saprophytic; onion skin bait; Sphagnum bog water; near Warsaw, Poland

Nowakowskiella multispora Karling; Sydowia, 17: 314-319, figs.1-8. 1964
Saprophytic; corn leaves and cellophane; soil, dry catch basin; Rhamnad Dist., India

Nowakowskiella pitcairnensis Karling; Nova Hedwigia, 15: 191-201, figs. 1-19. 1968
Saprophytic; hemp seed bait; soil; Pitcairn, Oceania

Nowakowskiella profusa Karling 1944 = Nowakowskiella elegans (Nowakowski) Schroeter
1893; See T.W. Johnson; Mycologia, 69: 34-45. 1977

Nowakowskiella profusa forma constricta Kobayasi & Konno; Bull. Natl. Sci. Mus., 14:
373-386, fig. 3 A-B. 1971. Saprophytic; cellophane; soil; Papua, Territory of New Guinea
and Solomon Islands

Nowakowskiella sculptura Karling; Trans. Brit. Mycol. Soc., 44: 453-457, figs. 1-24. 1961
Saprophytic; cellophane bait; soil; Baton Rouge, LA, USA.

Obelidium megarhizum Willoughby; Trans. Brit. Mycol. Soc., 44: 586-592, fig. 1, Pl. 37. 1961
Saprophytic; termite wings; submerged mud; Malham Tarn, U.K.

163

OEDOGONIOMYCETACEAE D.J.S. Barr [M]; HP, p. 463. 1990

Oedogoniomyces from Harpochytriales to Monoblepharidales; D.J.S. Barr: HP, p. 463. 1990

Olpidium (Braun) Rabenhorst 1868 = Pleotrachelus Zopf 1884; Karling: CI, p. 14. 1977
D.J.S. Barr placed Olpidium in Spizellomycetales; Canad. J. Bot., 58: 2380-2394. 1980

Olpidium appendiculatum Karling; Sydowia, 19: 213-226, Pl. 46, figs. 1-13. 1965
Saprophytic; pollen; sand & moss; near glacier, Antarctica

Olpidium bornovanus (Sahtiyanci) Karling; CI, p. 13. 1977; = Pleotrachelus bornovanus
Sahtiyanci 1962; = O. radicale; Lange & Insunza; Trans. Brit. Mycol. Soc. 69: 377-384. 1977
= O. cucurbitacearum Barr & Dias; Campbell & Sim; Canad. J. Bot., 72: 1136-1143. 1994

Olpidium brassicae (Woronin) Dangeard = Pleotrachelus brassicae (Woronin) Sahtiyanci;,
Arch. Mikrobiol., 41: 187-228. 1962

Olpidium cucurbitacearum D.J.S. Barr and Dias; Canad. J. Bot., 46: 1087-1091, figs. 1-12 &
13-22. 1968. = Olpidium radicale as redescribed, Lange & Insunza; Trans. Brit. Mycol. Soc.,
69: 377-384. 1977. = O. bornovanus (Sahiyanci) Karling 1962; see Campbell & Sim;
Canad. J. Bot., 72: 1136-1143. 1994. Parasitic; cucumber roots; soil; southern Ontario, Canada

Olpidium entophlyctoides in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum

Olpidium fulgens (Zopf) Karling; CI, p. 18. 1977; = Pleotrachelus fulgens Zopf 1892

Olpidium incognitum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, figs. 5-6. 1988
Parasitic; egg of rotifer; tank sediment; Lucknow U., India

Olpidium indicum Karling (homonym); Sydowia, 17: 302-307, figs. 1-21. 1964
= Olpidium indum Karling 1966
Parasitic; Catenophlyctis variabilis & Rhizophlyctis fuscus; soil; Rhamnad Dist., India

Olpidium indicum Kiran & Dyal (homonym & type not indicated); Proc. Natl. Acad. Sci, India,
Sect. B. 62: 295-298, fig. 2. 1992
Parasitic; root hairs of angiosperms; pond; Ramkatora, Varanasi, U.P., India

Olpidium indum Karling; Sydowia, Beiheft VI, p.6. 1966; = Olpidium indicum Karling 1964

Olpidium longum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, figs. 2-4. 1988
Parasitic; eggs of animicules; tank sediment; Lucknow U., India

Olpidium nematodae Skvortzow; Tribe (Trans. Brit. Mycol. Soc., 69: 509-511. 1977) refutes
Sparrow 1960 interpretation.

Olpidium poreferum Kiran and Dyal (invalid; type not designated); Proc. Natl. Acad. Sci.,
India, Sect. B, 62: 295-298, PI. 1, fig. 1. 1992.
Parasitic; rotifers; pond, Agricultural Farm; B.H.U., U.P., India

Olpidium pseudoeuglenae S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 7.
1988. Parasitic; chytrid on Closterium; tank sediment; Lucknow U., India

Olpidium radicale Schwartz & Cook; redescribed by Lange & Insunza; Trans. Brit. Mycol. Soc.,
69: 377-384. 1977. =O. cucurbitacearum D.J.S. Barr and Diaz 1968; Campbell & Sim (Canad.
J. Bot. 72: 1136-1143. 1994) consider O. radicale a nomen dubium and accept O. bornovanus
(Sahtiyanci) Karling.

Olpidium rostriferum var. indica Kiran & Dayal (invalid; type not designated); Proc. Natl.
Acad. Sci., India, Sect. B, 62: 295-298, fig. 3. 1992
Saprophytic; leaf of angiosperm; pond; Ramkatora, U.P., India

Olpidium sparrowii 1.J. Dogma; & table of Olpidium spp. parasitic on rotifers; Philipp. J. Biol.,
6: 9-20, figs. 1-16. 1977; Parasitic; rotifer eggs; cultivated soils; Putiao, Luzon, Philippines

Olpidium synchytrii Karling; Mycologia, 50: 944-947, figs. 1-7. 1958
Parasitic; Synchytrium namae on Nama jamaicensis plants; Texas, USA

Olpidium vermicola G.L. Barron & E. Szijarto; Mycologia, 78: 972-975, figs. 1-12. 1986
Parasitic; nematode eggs; old stump; Lake Manitowabing, Ontario, Canada

Olpidium virulentus (Sahtiyanci) Karling; CI, p. 13, 1977 = Pleotrachelus virulentus Sahtiyanci
1962

Olpidium wildemani (Petersen) Karling; CI, p. 18, 1977 = Pleotrachelus wildemani Petersen
1910

Olpidium zopfianus (Morini) Karling; CI, p. 18, 1977; = Pleotrachelus zopfianus Morini 1913

164

ORPINOMYCES D.J.S. Barr & H. Kudo [N]; Canad. J. Bot., 67: 2815-2824. 1989
Emended by Y.W. Ho & D.J.S. Barr; Mycologia, 87: 656-678. 1995

Orpinomyces bovis D.J.S. Barr & H. Kudo; Canad. J. Bot., 67: 2815-2824. figs. 37-51. 1989
= Orpinomyces joyonii (Breton et al.) Li et al. 1991
Saprophytic; plant material; digestive tract of herbivores; Lethbridge, Alberta, Canada

Orpinomyces intercalaris Y.W. Ho; Mycotaxon, 50: 139-150, figs. 1-10. 1994
Saprophytic; guinea grass (Panicum maximum), rumen of cattle (Bos indicus); Malaysia

Orpinomyces joyonii (Breton et al.) Li et al, Canad. J. Bot., 69: 580-589. 1991
= O. bovis Barr & H. Kudo 1989; = Neocallimastix joyonii Breton et al. 1989
Lectotype: see Y.W. Ho & D.J.S. Barr; Mycologia, 87: 656-678. 1995

Phlyctidium (Braun) Rabenhorst 1868 = Rhizophydium Schenk 1858
Karling; CI, Pp. 64-65. 1977

Phlyctidium anatropum (Braun) Rabenhorst 1856 = Rhizophydium anatropum (Braun) Karling;
CE p203. 197%

Phlyctidium apophysatum Canter 1947 emended by H. Canter; Trans. Brit. Mycol. Soc., 44:
522-528, figs. 1-3 & Pl. 34, figs.1-10. 1961. = Canteria apophysata (Canter) Karling 1971
Parasitic; Mougeotia; E.L.D., U.K.

Phlyctidium brevipes var. marinum Kobayasi & Ookubo 1954 = Rhizophydium brevipes var.
marinum (Kobayasi & Ookubo) Karling; CI, p. 65, 1977

Phlyctidium bumilleriae Couch 1932 = Rhizophydium bumilleriae (Couch) Karling; CI, p. 65.
1977

Phlyctidium globosum Skuja 1956 = Rhizophydium skujai (Skuja) Karling; CI, p. 64. 1977

Phlyctidium keratinophilum Ookubo and Kobayasi 1955 = Rhizophydium ellipsoidium (Ookubo
& Kobayasi) Karling; CI, p. 64. 1977

Phlyctidium keratinophilum var. savulescui Cejp; Omagiu lui Traian Savulescu (Bucuresti),
129-138, fig. 1. 1959. Saprophytic; human hair; rill; pasture near Rokycany, Romania

Phlyctidium marinum Karling; Sydowia, 20: 74-84, Pl. XII, figs. 1-8. 1967
= Rhizophydium novae-zelandiensis (Karling) Karling; CI, p. 65. 1977
Saprophytic; Pinus pollen; beach sand + sea water; Aukland, N.Z.

Phlyctidium mycetophagum Karling 1946 = Rhizophydium obypriformis (Karling) Karling; CI,
p. 64. 1977

Phlyctidium olla Sparrow 1933; = Rhizophydium olla (Sparrow) Karling; CI, p. 65. 1977

Phlyctidium scenedesmi Fott; Z. Allg. Mikrobiol., 7: 97-102, figs. 1-16. 1967
= Rhizophydium scenedesmi (Fott) Karling; CI, p. 65. 1977
Parasitic; Scenedesmus quadricauda in mass culture; water; Trebon, Czechoslovakia

Phlyctidium spinulosum Sparrow 1933 = Rhizophydium spinulosum (Sparrow) Karling; CI,
p65, 1977

Phlyctidium tenue Sparrow 1952 = Rhizophydium tenue (Sparrow) Karling; CI, p. 65. 1977

Phlyctidium tubulatum in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum

Phlyctochytrium acuminatum D.J.S. Barr; Canad. J. Bot., 47: 991-997, figs. 71-82. 1969
= Spizellomyces acuminatus (D.J.S. Barr) D.J.S. Barr 1984
Saprophytic; pine pollen; soil; southwestern Ontario, Canada

Phlyctochytrium aestuarii Ulken, Veroff. Inst. Meeresforsch. Bremerhaven, 13: 205-216, figs. 5
& 6. 1972 = Rhizophydium aestuarii (Ulken) Amon 1984
Saprophytic; pine pollen; river mud; Weser Estuary, Germany

Phlyctochytrium africanum Gaertner 1954 (not validly published) = Spizellomyces acuminatus
(D.J.S. Barr) D.J.S. Barr; Canad. J. Bot., 62: 1171-1201. 1984

Phlyctochytrium apophysatum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig.
36. 1988. Saprophytic ?; Closterium; tank; garden, Lucknow U., India

Phlyctochytrium arcticum D.J.S. Barr; Canad. J. Bot., 48: 2279-2283, figs.1-17. 1970
= Triparticalcar arcticum D.J.S. Barr 1984
Saprophytic; pollen; soil, pH 7.9; Canadian Arctic

165

Phlyctochytrium aureliae Ajello 1945 = Phlyctochytrium variable 1954; T.W. Johnson; Norw.
J. Bot., 22: 249-257. 1975

Phlyctochytrium californicum D.J.S. Barr; Canad. J. Bot., 47: 991-997, figs. 23-33. 1969
= Gaertneriomyces semiglobiferus (Uebelmesser) D.J.S. Barr 1980
Saprophytic; pine pollen; soil; CA, USA

Phlyctochytrium chandleri Sparrow (IF 4:385); Arch. Microbiol., 104: 39-49, figs. 6-9. 1975
Saprophytic; pollen; soil; oasis, Tunisia

Phlyctochytrium circulidentatum Koch; In Umphlett & Koch; Mycologia, 61: 1021-1030, figs.
1-3, 11-17. 1969. Saprophytic; pollen and moribund Closterium; bottom debris; Ravenal
Lake, NC, USA

Phlyctochytrium cystoferum Willoughby; Arch. Mikrobiol., 52: 101-131, fig. 1 a-h. 1965
= Polyphlyctis cystofera (Willoughby) A. Batko; ZH, p. 183. 1975
Saprophytic; grass leaf bait; soil from heath; Victoria, Australia

Phlyctochytrium dichotomum Umphlett; Mycologia, 59: 1085-1096, figs. 1-41. 1967
= Kochiomyces dichotomus (Umphlett) D.J.S. Barr 1980
Saprophytic; sweet gum pollen; soil + sawdust; Chatham Co., NC, USA

Phlyctochytrium dissolutum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 37.
1988. Saprophytic ?; Closterium; tank; garden, Lucknow U., India

Phlyctochytrium furcatum Sparrow; Arch. Mikrobiol., 53: 178-180, figs. 1-9. 1966
Saprophytic; pine pollen; almost dry bog with Vaccinium; MI, USA

Phlyctochytrium hirsutum Karling; Sydowia, 20: 86-95, figs. 3-10. 1967
Saprophytic; purified shrimp chitin; soil; N.Z.

Phlyctochytrium incrustans Sparrow & Lange; Canad. J. Bot., 55: 1879-1890, figs. 19-22. 1977
Saprophytic; pine pollen; bog; Emmet Co., MI, USA

Phlyctochytrium indicum Karling; Sydowia, 17: 285-296, figs. 16-31. 1964
Saprophytic; pine pollen; brackish soil; Rhamnad District, Madras, India

Phlyctochytrium irregulare Koch; J. Elisha Mitchell. Sci. Soc., 73: 108-122, figs. 13-24. 1957
Saprophytic; Liguidambar pollen and Paspalum leaves bait; wet soils; NC, USA

Phlyctochytrium kniepii Gaertner 1954 (not validly published) = Spizellomyces kniepii Gaertner
ex D.J.S. Barr; Canad. J. Bot., 62: 1171-1201. 1984

Phlyctochytrium lackeyi Sparrow; Mycologia, 68: 1233-1234, figs. 1-4. 1976
Saprophytic; pollen; Punch Bowl; Milford, FL, USA

Phlyctochytrium macrosporum Sparrow; Mycologia, 70: 140-149, figs. 1-14. 1978
Parasitic?; live pine pollen; neuston; Newnan's Lake, FL, USA

Phlyctochytrium mangrovii Ulken; Veroff. Inst. Meeresforsch. Bremerhaven, 13: 217-230, figs.
1, 2. 1972. Saprophytic; pine pollen & cattle hair; mangrove muds; Cananeia, Sao Paulo, Brazil

Phlyctochytrium marilandicum Sparrow; Mycologia, 65: 1331-1336, figs. 1-11. 1973
Saprophytic; pollen; woodland soil; MD, USA

Phlyctochytrium megastomum Karling; Mycopathol. Mycol. Appl., 36: 165-178, figs. 1-17.
1968 =P. irregularis Koch 1957 (Sparrow opinion, unpublished)
Saprophytic; bleached corn leaves; soil; Rarotonga, Cook Islands

Phlyctochytrium mucosum Sparrow & I.J. Dogma; Arch. Mikrobiol., 89: 177-204, fig. 1, J-M.
1973. Saprophytic; pine pollen bait; soil; Dominican Republic

Phlyctochytrium multidentatum Umphlett; Mycologia, 61: 1021-1030, figs. 4-10. 1969
Saprophytic; sweet gum pollen; soil & water; Chapel Hill, NC, USA

Phlyctochytrium neuhausiae Sparrow (IF 4:417) ; Proc. lowa Acad. Sci., 81: 2-5, figs. 10-14.
1974. Saprophytic; pine pollen; soil; Delphi, Greece

Phlyctochytrium palustre Gaertner 1954 (not validly published) = Spizellomyces palustris
Gaertner ex D.J.S. Barr; Canad. J. Bot. 62: 1171-1201. 1984

Phlyctochytrium parasitans Sparrow & 1.J. Dogma; Arch. Mikrobiol., 89: 177-204, fig. 2, T-U.
1973. Parasitic; eggs of Achlya flagellata; damp soil; Carretara Duarte, Hispaniola

Phlyctochytrium peruvianum Knox (not validly published) PhD thesis, VPI, Blacksburg, VA.
1970. = Spizellomyces punctatus (Koch) D.J.S. Barr 1984

166

Phlyctochytrium planicorne Atkinson 1909 emended by Umphlett & Holland; Mycologia, 52:
429-435. 1960; = Rhizophydium digitatum Scherffel 1926; T.W. Johnson: Norw. J. Bot., 22:
249-257. 1975

Phlyctochytrium plurigibbosum D.J.S. Barr, Canad. J. Bot., 47: 991-997, figs. 44-52. 1969
= Spizellomyces plurigibbosus (D.J.S. Barr) D.J.S. Barr 1984
Saprophytic; pine pollen; soil; Ottawa, Ontario, Canada

Phlyctochytrium powhatanense Roane (IF 4:250); Mycologia, 65: 531-538, figs. 1-2, 15-20.
1973. Saprophytic; rotifer, chitin, snake skin; impoundment; Pulaski Co., VA, USA

Phlyctochytrium punctatum Koch; J. Elisha Mitchell. Sci. Soc., 73: 108-122, figs. 1-16. 1957
= Spizellomyces punctatus (Koch) Barr 1984
Saprophytic; Liqguidambar and pine pollen bait; soil; New Guinea

Phlyctochytrium recurvastomum Knox & Paterson; Mycologia, 65: 373-387, figs. 11-18. 1973
Saprophytic; pollen; soil; McMurdo area, Antarctica

Phlyctochytrium reinboldtae Persiel; Arch. Mikrobiol. 32: 411-415, fig. 1. 1959

Saprophytic; pine pollen; soil ; Botanical Garden, Gottingen, Germany
Phlyctochytrium semiglobiferum Uebelmesser 1956 = Gaertneriomyces semiglobiferus

(Uebelmesser) D.J.S. Barr; Canad. J. Bot., 58: 2380-2394. 1980
Phlyctochytrium spinosum Sparrow; Mycologia, 65: 1331-1336, figs. 12-16. 1973

Saprophytic; pollen ?; soil from spring-fed meadow; HI, USA
Phlyctochytrium variable Rieth 1954; = Phlyctochytrium aureliae Ajello 1945; see

T.W. Johnson; Norw. J. Bot., 22: 249-257. 1975
Phlyctochytrium vaucheriae Rieth 1956 = Blyttiomyces vaucheriae (Rieth) I.J. Dogma;

Mycologia, 61: 1149-1158. 1969
Phlyctochytrium verruculosum Knox (not valid pub.) PhD thesis, VPI, Blacksburg, VA. 1970

= Spizellomyces punctatus (Koch) D.J.S. Barr ; Canad. J. Bot., 58: 2380-2394. 1980
Phlyctorhiza variabilis Karling 1947 = Entophlyctis variabilis (Karling) Cejp; Omagiu lui

Traian Savulescu, 129-138. 1959; = Catenophlyctis variabilis (Karling) Karling; Amer. J. Bot.,

52: 133-138. 1965
Physodermataceae; transferred from Chytridiales to Blastocladiales by Lange & Olson; Trans.

Brit. Mycol. Soc., 74: 449-457. 1980
PIROMYCES Gold, Heath & Bouchop [N]; Biosystems, 21: 403-415. 1988 = Piromonas

communis sensu Orpin; J. Gen. Microbiol., 99: 107-117. 1977
Piromyces communis Gold, Heath & Bauchop; BioSystems, 21: 403-415, (figs. 1-10 in Orpin; J.

Gen. Microbiol., 99: 107-117. 1977). 1988 = Piromonas communis sensu Orpin 1977

Saprophytic; plant material; rumen of sheep
Piromyces dumbonicus J. Li (IF); Canad. J. Bot., 68: 1021-1033, figs. 2, 4, 9, 12, 13, 19, 23, 24,

31. 1990; Saprophytic; cellulose; gut of elephant; New Zealand
Piromyces mae J. Li; Canad. J. Bot., 68: 1021-1033, figs. 1, 3, 5-8, 10, 11, 15-16, 20-22, 25-30,

32. 1990. Saprophytic; plant material; caecum of horse; New Zealand
Piromyces minutus Y.W. Ho; Mycotaxon, 47: 285-293, figs. 1-14. 1993

Saprophytic; guinea grass (Panicum maximum); rumen of Sika deer (Cervus nippon); Malaysia
Piromyces rhizinflatus Breton et al. (IF 6:445); FEMS Microbiol. Lett., 82: 1-8, figs. 1-2. 1991

Lectotype: see Y.W. Ho & D.J.S. Barr; Mycologia, 87: 656-678. 1995

Saprophytic; culture; dried faeces of Saharian ass; Tunisia
Piromyces spiralis Y.W. Ho; Mycotaxon, 48: 59-68, figs. 1-14. 1993.

Saprophytic; grass (Panicum maximum); rumen of katjang goat (Capra hircus); Malaysia
Pleotrachelus Zopf 1884; = Olpidium (Braun) Rabenhorst 1868; Karling, CI, p. 14. 1977
Pleotrachelus askaulos Bradley (fossil); Amer. J. Bot., 54: 577-582, fig. 8. 1967

Eocene Green River Formation; Wyoming, USA

167

Pleotrachelus bornovanus Sahtiyanci; Arch. Mikrobiol., 41: 187-228, figs. 50-79. 1962.
= Olpidium bornovanus (Sahtiyanci) Karling; CI, p. 13. 1977; = O. radicale (sensu Lange &
Insunza 1977); = O. cucurbitacearum D.J.S. Barr & Dias 1968; See Campbell & Sim; Canad. J.
Bot., 72: 1136-1143. 1994. Parasitic; Lactuca, Spinacea, Beta, Nicotiana, Solanum et al.; soil

Pleotrachelus brassicae (Woronin) Sahtiyanci; Arch. Mikrobiol. 41: 187-228. 1962 = Olpidium
brassicae (Woronin) Dangeard; maintained as O. brassicae by Karling; CI, p. 18. 1977

Pleotrachelus virulentus Sahtiyanci; Arch. Mikrobiol., 41: 187-228, figs. 24-49. 1962
= Olpidium virulentus (Sahtiyanci) Karling; CI, p. 13. 1977
Parasitic; Lactuca, Solanum, Beta, Nicotiana, Capsella

Pleotrachelus wildemanni Petersen 1910 = Olpidium wildemanni (Petersen) Karling; CI, p. 18.
1977

Pleotrachelus zopfianus Morini 1913 = Olpidium zopfianus (Morini) Karling; CI p. 18. 1977

Podochytrium chitinophilum Willoughby; Trans. Brit. Mycol. Soc., 44: 586-592, fig. 2, Pl. 38.
1961. Saprophytic; termite wings; Wastwater; E.L.D., U.K.

Podochytrium dentatum Longcore; Mycologia, 84: 183-192, figs. 1-29. 1992
Saprophytic; chitin; lake, circumneutral pH; Maine, USA

Podochytrium ellerbeckense Willoughby; Nova Hedwigia, 5: 335-340, Pl. 53 a-w. 1963
Saprophytic; waterlily leaf stalks; garden pond; E.L.D., U.K.

Polyphagus asymmetricus Valkanov (no Latin diagnosis); Arch. Protistenk., 106: 565-568, PI.
44. 1963. Parasitic; Botrydiopsis; hyponeuston; southern Bulgaria

Polyphagus elegans Canter; Nova Hedwigia, 5: 419-428, Pls. 74-76. 1963
Parasitic; Ulothrix mucosum; plankton; Windermere, E.L.D., U.K.

Polyphagus hyponeustonica (Valkanov) Karling; CI, p. 174. 1977 (sp. not validated)
= Arnaudovia hyponeustonica Valkanov 1963 (no Latin diagnosis)

Polyphagus serpentinus Canter; Nova Hedwigia, 5: 419-428, Pls. 71-73. 1963
Parasitic; Spondylosium spp.; lakes; E.L.D., U.K.

Polyphagus starrii Johns; Mycologia, 56: 441-451, figs. 1-6. 1964
Parasitic; Eudorina, Pandorina, et al.; Stonewall Pond; MA, USA

POLYPHLYCTIS Karling [C]; Sydowia, 20: 86-95. 1967

Polyphlyctis cystofera (Willoughby) A. Batko; ZH, p. 183. 1975 = Phlyctochytrium cystoferum
Willoughby 1965

Polyphlyctis unispina (Paterson) Karling; Sydowia, 20: 86-95. 1967 = Phlyctochytrium
unispinum Paterson 1956

POWELLOMYCES Longcore, D.J.S. Barr & N. Désaulniers [S]; Canad. J. Bot. 73: 1385-1390.
1995

Powellomyces hirtus Longcore, D.J.S. Barr & N. Désaulniers; Canad J. Bot. 73: 1385-1390, figs.
1-18. 1995. Saprophytic; pollen bait; soil; Ontario, Canada

Powellomyces variabilis Powell & Koch ex Longcore, D.J.S. Barr & N. Désaulniers; Canad. J.
Bot. 73: 1385-1390. 1995. = Entophlyctis variabilis Powell & Koch 1977 not E. variabilis
(Karling) Cejp 1959

Pringsheimiella dioica Couch 1939 (homonym and no Latin diagnosis) = Dictyomorpha dioica
Couch ex Mullins; Amer. J. Bot., 48: 377-387. 1961

PSEUDOPILEUM Canter [C]; Trans. Brit. Mycol. Soc., 46: 305-320. 1963

Pseudopileum unum Canter; Trans. Brit. Mycol. Soc., 46: 305-320, fig. 3. 1963
Parasitic; cysts of Mallomonas sp.; plankton of Elterwater; E.L.D., U.K.

Rhizidium endosporangiatum Karling; Mycopathol. Mycol. Appl., 36: 165-178, figs. 18-37.
1968. Saprophytic; bleached corn leaves & pollen; soil sample; Pitcairn & Niue, Oceania

Rhizidium megastomum Sparrow; Mycopathol. Mycol. Appl., 25: 119-143, figs. 19-23. 1965
Saprophytic; snake skin; soil; Oahu, HI, USA

Rhizidium microcystidis Sen; Arch. Hydrobiol. (Suppl.), 79: 177-184, figs. 1, 2. 1988
Parasitic; Microcystis aeruginosa (Cyanobacteria); Shearwater; Wiltshire, U.K.

Rhizidium parasiticum in S.N. Dasgupta; Indian Phytopathol., 35: 215. 1982; ? nomen nudum

168

Rhizidium renifore Karling (IF 4:43); Arch. Mikrobiol., 70: 266-287, fig. 1 A-O. 1970
Saprophytic; purified shrimp and lobster chitin; soil sample; Auckland Province, N.Z.

Rhizidium tomiyamanum Konno; J. Jap. Bot., 44: 315-317, figs. A-H. 1969
Saprophytic; Porphyra, pine pollen bait; marine; ? Japan

Rhizoclosmatium hyalinum Karling; Sydowia, 20: 96-108, figs. 48-59. 1967
Saprophytic; exuviae and chitin bait; water in outdoor tub; U. of Otago, N.Z.

Rhizophlyctis A. Fischer 1892 sensu Sparrow 1960; I.J. Dogma designated R. mastigotrichis
(Nowakowski) Fischer as lectotype; Nova Hedwigia, 24: 393-411. 1973; but Dogma’s lectotype
was predated by designation of R. rosea (De Bary & Woronin) Fischer in Clements & Shear;
The Genera of Fungi, p. 235. 1931.

Rhizophlyctis aurantiaca 1.J. Dogma; Nova Hedwigia, 25: 51-89, Pls. I-III. 1974
Saprophytic; chitin; soil; Annapolis, MD, USA

Rhizophlyctis boninensis Kobayasi & Konno; Bull. Natl. Sci. Mus., 12: 725-733, fig. 3 L-Q.

1969. Saprophytic; ? cellophane; soil; Chichijima, Bonin Islands

Rhizophlyctis bonseyi Sparrow; Mycopathol. Mycol. Appl., 25: 119-143, figs. 29-54. 1965
Saphrophytic; chitin; soil, margin of reservoir; Maui, HI, USA

Rhizophlyctis columellae S.N. Dasgupta & R. John; Indian Phytopathol., 43: 564-567, fig. 1.
1990 (1991). Parasitic; zygospore of Spirogyra sp.; tank; Lucknow U., India

Rhizophlyctis costatus Konno; Sci. Rep. Tokyo Kyoiku Daigaku Sect. B, 14: 227-292, Pl. 8
A-D. 1972. Saprophytic; cicada wings; forest soil; Ohdaigahara (Nara Prefecture), Japan

Rhizophlyctis fuscus Karling (IF 3:310); Mycopathol. Mycol. Appl., 23: 215-222, figs. 12-21.
1964. Saprophytic; human & snake skin, corn leaf; soil; Madras, India

Rhizophlyctis hirsutus Karling; Mycopathol. Mycol. Appl., 23: 215-222, figs. 22-26. 1964
Saprophytic; skin, corn leaves, hemp seed; soil; dry rice paddy, Madras, India

Rhizophlyctis ingoldii Sparrow; Trans. Brit. Mycol. Soc., 40: 523-535, fig. 1 F-K. 1957
Saprophytic; chitin bait; soil; widespread, U.K. (in Sparrow 1960 as footnote)

Rhizophlyctis lovettii Karling (IF 3:310); Mycopathol. Mycol. Appl., 23: 215-222, figs. 1-11.
1964. Saprophytic; human fibrin film; soil; Madras, India

Rhizophlyctis oceanis Karling; Amer. J. Bot., 56: 211-221 figs. 1-108. 1969
Saprophytic; purified shrimp chitin; soil; Pitcairn et al. Islands, Oceania

Rhizophlyctis oceanis var. floridaensis Karling; Nova Hedwigia, 28: 209-229, figs. 17-28. 1976
Saprophytic; chitin; soil, flower garden; FL, USA

Rhizophlyctis petersenii var. appendiculata Karling; Sydowia, 20: 96-108, figs. 60-67. 1967
Saprophytic; corn leaves and chitin; soil and water; N.Z.

Rhizophlyctis reynoldsii 1.J. Dogma; Nova Hedwigia, 25: 51-89, Pl. VII., figs. 1-19. 1974
Saprophytic; chitin; leaf litter; Rangoon, Burma

Rhizophlyctis serpentina 1.J. Dogma (IF 4:318); Nova Hedwigia, 25: 51-89, Pl. [V-VI, figs.
1-45. 1974. Saprophytic; chitin bait; soil & leaf litter; widespread, Cameroon

Rhizophlyctis tropicalis Sparrow & 1.J. Dogma; Arch. Mikrobiol., 89: 177-204, fig. 4, A-M.
1973. Saprophytic; lens paper and onion skin baits; soil; Dominican Republic

Rhizophlyctis variabilis Karling; Sydowia, 20: 96-108, figs. 68-81. 1967
Saprophytic; bleached corn leaves; soil; N.Z.

Rhizophlyctis variabilis var. burmaensis 1.J. Dogma; Nova Hedwigia, 25: 51-89, Pl. VIII, figs.
1-20. 1974. Saprophytic; chitin; soil & leaf litter; Burma

Rhizophlyctis willoughbyi Konno; Sci. Rept. Tokyo Kyoiku Daigaku Sect. B, 14: 227-292. Pl. 7
A-D. 1972. Saprophytic; bee wings, cicada wings, & snake skin; soils; widespread, Japan

Rhizophydium Schenk; all Phlyctidium spp. into Rhizophydium; Karling; CI, p. 64. 1977

Rhizophydium achnanthis Friedmann 1952 = Hapalopera achnanthis (Friedmann) A. Batko;
ZH, p. 170. 1975

Rhizophydium aestuarii (Ulken) Amon; Mycologia, 76: 132-139. 1984 = Phlyctochytrium
aestuarti Ulken 1972

Rhizophydium anatropum (Braun) Karling; CI, p. 65. 1977 = Phlyctidium anatropum Braun
1856

169

Rhizophydium androdioctes Canter; Trans. Brit. Mycol. Soc., 56: 115-120, figs. 1-19. 1971
Parasitic; Dictyosphaerium pulchellum (Green alga); Loch Leven; Kinross, Scotland

Rhizophydium angulosum Karling; Arch. Mikrobiol., 61: 112-127, fig. 1 M-X. 1968
Saprophytic; pollen and bleached corn leaves; many soil collections; Oceania

Rhizophydium annulatum Sparrow; Canad. J. Bot., 55: 1501-1504, figs.1-6. 1977
Saprophytic; pine pollen; Punch Bowl; Melrose, FL, USA

Rhizophydium aphanomycis Karling; Nova Hedwigia, 27: 747-771, figs. 12-21. 1976
Parasitic; Aphanomyces sp.; soil; Ngorongora Crater, Tanzania

Rhizophydium aureum Konno; Sci. Rep. Tokyo Kyoiku Daigaku Sect. B, 14: 227-293, P1.3,
figs. A-D. 1972. Saprophytic; snake skin, bee wings, & cicada wings; soil; Tsu-shima, Bonin
Island, Japan

Rhizophydium biporosum (Couch) D.J.S. Barr; Canad. J. Bot., 51: 967-975. 1973
= Phlyctochytrium biporosum Couch 1932; = Phlyctidium bumilleriae Couch 1932

Rhizophydium blastocladianum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig.
16. 1988. Parasitic; Blastocladia on apple; tank sediment; Lucknow U., India

Rhizophydium blyttiomycerum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig.
24. 1988. Parasitic; Blyttiomyces spinosus & zygospore of Spirogyra; ditch; Sultanpur Road,
SE of Lucknow U., India

Rhizophydium brevipes var. marinum (Kobayasi & Ookubo) Karling; 1977., p. 65
= Phlyctidium brevipes var..marinum Kobayasi & Ookubo 1954

Rhizophydium bumilleriae (Couch) Karling; CI, p. 65. 1977 = Phlyctidium bumilleriae Couch
1932; = Rhizophydium biporosum (Couch) D.J.S. Barr; Canad. J. Bot., 51: 967-975. 1973

Rhizophydium capillaceum D.J.S. Barr; Canad. J. Bot., 47: 991-997, figs. 12-22 on Pls. I-VI.
1969. Saprophytic; pine pollen; soil; Ottawa, Ontario, Canada

Rhizophydium clavatum Karling; Sydowia, 20: 74-84, figs. 21-24. 1967
Saprophytic; chitin bait; soil sample; N.Z.

Rhizophydium clavatum S.N. Dasgupta & R. John (homonym); Bull. Bot. Surv. India, 30: 1-82
fig. 34. 1988. Parasitic; Closterium; tank; Lucknow U., India

Rhizophydium coleochaetes (Nowakowski) Fischer 1892 = Chytridium coleochaetes
Nowakowski 1876; H. Canter places in Chytridium; Trans. Brit. Mycol. Soc. 43: 14-18. 1960

Rhizophydium collapsum Karling; Sydowia, 17: 285-296, figs. 1-15. 1964
Saprophytic; pine pollen; brackish soil; Rhamnad District, India

Rhizophydium conchiforme S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 35.
1988. Saprophytic ?; Spirogyra; ditch; Sultanpur Road, SE of Lucknow U., India

Rhizophydium condylosum Karling; Arch. Mikrobiol., 61: 112-127, fig. 2, A-X. 1968
Saprophytic, human hair and snake skin; soil; Cook Island, Fiji, & Pitcairn

Rhizophydium contractophilum Canter; Trans. Brit. Mycol. Soc., 42: 185-192, figs. 1-4. 1959
Parasitic; Eudorina elegans and Eudorina sp.; Lake Windermere; E.L.D., U.K.

Rhizophydium coralloidum Karling (IF 4:484); Mycologia, 68: 1245-1248, figs. 1-9. 1976
Parasitic; Karlingia rosea; soil; Canaveral FL, USA

Rhizophydium difficile Canter 1954 = Hapalopera difficilis (Canter) A. Batko; ZH, p. 169.
1975

Rhizophydium digitatum Scherffel 1926 = Phlyctochytrium planicorne Atkinson 1909; see
T.W. Johnson; Norw. J. Bot., 22: 249-257. 1975

Rhizophydium dubium Kiran & Dayal (invalid; type not designated); Indian Phytopathol., 44:
30-34, Pl. 1.1991. Saprophytic; rotting leaves of Eichornia; pond; Agricultural Farm,
Varanasi, India

Rhizophydium echinocystoides Sparrow; J. Elisha Mitchell Sci. Soc., 84: 62-68, figs. 15-18.
1968. Saprophytic; pine pollen; Sphagnum bog; Humbolt Co, CA, USA

Rhizophydium elyensis Sparrow; Trans. Brit. Mycol. Soc., 40: 523-535, fig 2 I-M. 1957
Saprophytic; snake skin bait; soil; Ely, Isle of Ely, U.K. (in Sparrow 1960 as footnote)

Rhizophydium ellipsoidium (Ookubo & Kobayasi) Karling; CI, p. 64. 1977 = Phlyctidium
keratinophilum Ookubo & Kobayasi 1955

bf]

170

Rhizophydium fragilariae Canter 1950 = Hapalopera fragilariae (Canter) A. Batko; ZH, p.
170. 1975

Rhizophydium fugax Canter; Trans. Brit. Mycol. Soc., 51: 699-705, Pl. 57 & fig. 1. 1968
Parasitic; Cryptomonas spp.; plankton of lakes; E.L.D., U.K.

Rhizophydium gonapodyanum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig.
23. 1988. Parasitic; sporangium of Gonapodya polymorpha, tank sediment; Lucknow U., India

Rhizophydium hispidulosum Seymour, J. Elisha Mitchell Sci. Soc., 86: 187-190. figs. 1-16.
1970. Saprophytic; cockroach wing bait; soil from ditch; Morehead City, NC, USA

Rhizophydium horizontale Paterson, Mycologia, 50: 85-96, fig. 1 H-K. 1958
Parasitic; Stephanodiscus sp.; plankton, Huron River & Portage Lake; Washtenaw Co. MI, USA

Rhizophydium karlingii Sparrow 1960 = R. laterale (Braun) Rabenhorst; D.J.S. Barr, Canad. J.
Bot., 51: 967-975. 1973

Rhizophydium lagenaria S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 21.
1988. Parasitic; sporangium of Blastocladia on fruit bait; tank sediment; Lucknow U., India

Rhizophydium laterale (Braun) Rabenhorst = R. karlingii Sparrow 1960; D.J.S. Barr, Canad. J.
Bot., 51: 967-975. 1973

Rhizophydium lenelangeae Sparrow (IF 4:484); Canad. J. Bot., 55: 1501-1504, figs. 7-15. 1977
Saprophytic; pine pollen bait; river bank soil; Guatemala

Rhizophydium littoreum Amon; Mycologia, 76: 132-139, figs. 1-12. 1984
?Saprophytic; Bryopsis plumosa, marine, Atlantic coast, USA

Rhizophydium macroporosum Karling; Sydowia, 20: 74-84, figs. 9-20. 1967
Saprophytic; snake skin and bleached corn leaves; soil samples; N.Z.

Rhizophydium manoense Sparrow; Mycopathol. Mycol. Appl., 25: 119-143. figs. 1-7. 1965
Parasitic; diatom (Cocconeis); stream; Oahu, HI, USA

Rhizophydium melosirae Friedman 1952 = Hapalopera melosirae (Friedman) A. Batko; ZH, p.
L705 1975

Rhizophydium mougeotiae Pongratz; Schweiz. Z. Hydrol., 28: 104-132, fig. 2 & Pl. 5, 33-36.
1966. Parasitic; Mougeotia gracillima; plankton; Lake Léman, Switzerland

Rhizophydium nobile Canter; Proc. Linn. Soc. London, 179: 197-201, figs. 1-8. 1968
Parasitic; resting spores of Ceratium hirundinella; lakes; E.L.D., U.K.

Rhizophydium novae-zeylandiensis (Karling) Karling; CI, p. 65. 1977 = Phlyctidium marinum
Karling 1967

Rhizophydium obypriformis (Karling) Karling; CI, p. 64. 1977 = Phlyctidium mycetophagum
Karling 1956

Rhizophydium olla (Sparrow) Karling; CI, p. 65. 1977 = Phlyctidium olla Sparrow 1933

Rhizophydium patellarium Scholz; Arch. Mikrobiol., 29: 354-362, fig. 2. 1958
Saprophytic; pine pollen bait; soils; northern Spain and Germany

Rhizophydium pedicellatum Paterson 1956; emended by T.W. Johnson; Norw. J. Bot., 23:
139-151. 1976

Rhizophydium pelagicum Paterson; Mycologia, 50: 85-96, fig. 1 A-G. 1958
Parasitic; Colacium sp; plankton, Frains Lake; Washtenaw Co., MI, USA

Rhizophydium piriformis (Fott) Karling; CI, p. 65. 1977 = Hapalopera piriformis Fott 1942

Rhizophydium planktonicum Canter 1948; emended by Canter; J. Linn. Soc., Bot., 62: 267-278,
fig. 1. 1969. Parasitic; Asterionella formosa (diatom); lakes; E.L.D., U.K.

Rhizophydium poculiforme S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, figs.
28-30. 1988. Parasitic; Closterium; tank; Lucknow U., India

Rhizophydium polystomum Karling; Sydowia, 20: 74-84, figs. 26-29. 1967 (Pls. XII-XIV)
Saprophytic; bleached corn leaves; soil samples; N.Z.

Rhizophydium porosum Sparrow & Lange; Canad. J. Bot., 55: 1879-1890, figs. 1-5. 1977
Saprophytic; pine pollen bait; Johnson Road Bog; Cheboygan Co., MI, USA

Rhizophydium proliferum Knox & Paterson; Mycologia, 65: 373-387, figs. 6-10. 1973
Saprophytic; pine pollen; soil or fresh water? Ross Island, Antarctica

171

Rhizophydium punctatum O.G. Golubeva; Novosti Sistematiki Nizshikh Rastenii, 25: 81-86, fig.
1. 1988. Saprophytic; pine pollen; Lake Teletzkoje; Altiaca Province, Russia

Rhizophydium rarotonganensis Karling; Arch. Mikrobiol. 61: 112-127, fig. 1 A-L. 1968
Parasitic; Nowakowskiella profusa; soil; Rarotonga, Cook Islands, Oceania

Rhizophydium reflexum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 18. 1988
Parasitic; sporangium of Blastocladia on Pyrus fruit; tank sediment; Lucknow U., India

Rhizophydium rhizinum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 17. 1988
Parasitic; sporangium of Blastocladia on Pyrus fruit; tank sediment; Lucknow U., India

Rhizophydium rotundum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, figs. 25-27.
1988. Parasitic; Closterium, Oedogonium, and Zygnema; tank; garden, Lucknow U., India

Rhizophydium scenedesmi (Fott) Karling; CI, p. 65. 1977 = Phlyctidium scenedesmi Fott 1967

Rhizophydium sibyllinum Sparrow (IF 4:420); Proc. Iowa Acad. Sci., 81: 2-5, figs. 1-9. 1974
Saprophytic; pine pollen; soil; Delphi, Greece

Rhizophydium signyense Willoughby; Nova Hedwigia, 22: 469-487, Pl. 2 m-t; 4 e, f. 1971
Saprophytic; chitin bait; soil; Signy Island, Antarctica

Rhizophydium skujai (Skuja) Karling; CI, p. 64. 1977 = Phlyctidium globosum Skuja 1948

Rhizophydium sparrowii S.N. Dasgupta & R. John; Indian Phytopathol., 43: 564-567, fig. 2.
1990 (1991). "Saproparasitic"; Ulothrix sp.; ditch; Sultanpur Rd, SE of Lucknow U., India

Rhizophydium sphaerocarpum var. rhizoclonii Huth & Gaertner; Trans. Brit. Mycol. Soc., 61:
431-434, Pl. 43. 1973. Parasitic; Rhizoclonium hieroglyphicum; brackish water; Bremerhaven,
Germany

Rhizophydium sphaerocarpum var. spirogyrae D.J.S. Barr; Canad. J. Bot., 48: 1607-1071, figs.
1-6. 1970. Parasitic; moribund Spirogyra; southwestern Ontario, Canada

Rhizophydium sphaerotheca Zopf 1887
Neotype: D.J.S. Barr; Canad. J. Bot., 47: 991-997, figs. 1-11. 1969

Rhizophydium spinosum Willoughby; Arch. Mikrobiol., 52: 101-131, fig. 1 i-w. 1965
Saprophytic; cellophane bait; soil; Victoria, Australia

Rhizophydium spinosum Karling (homonym); Nova Hedwigia, 35: 107-116, figs. 1-6, 1981
Saprophytic; cellophane & bleached corn leaves; soil samples; Sri Lanka

Rhizophydium spinosum S.N. Dasgupta & R. John (homonym); Bull. Bot. Surv. India, 30: 1-82,
fig. 20. 1988. Parasitic; Blastocladia on fruit bait; tank sediment; Lucknow U., India

Rhizophydium spinulosum (Sparrow) Karling; CI, p. 65. 1977 = Phlyctidium spinulosum
Sparrow 1933

Rhizophydium squamosum O.G. Golubeva; Novosti Sistematiki Nizshikh Rastenii, 25: 81-86,
fig. 2. 1988. Saprophytic; pine pollen; Lake Teletzkoje; Altiaca Province, Russia

Rhizophydium stellatum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 22. 1988
Parasitic; Sporangium of Gonapodya polymorpha; tank sediment; Lucknow U., India

Rhizophydium stipitatum Sparrow; Trans. Brit. Mycol. Soc., 40: 523-535, fig. 1 A-C. 1957
Saprophytic; snake skin bait; Cavenham Heath; Suffolk, U.K. (In Sparrow 1960 as footnote)

Rhizophydium tenue (Sparrow) Karling; CI, p. 65. 1977 = Phlyctidium tenue Sparrow 1952

Rhizophydium tetragenum Pongratz; Schweiz. Z. Hydrol., 28: 104-132, fig. 1 & Pl. 4, 25-28.
1966. Parasitic; Asterionella formosa; plankton; Lake Léman, Switzerland

Rhizophydium tubulatum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 31-33.
1988. Parasitic; Closterium; tank; Lucknow U., India

Rhizophydium ubiquetum Canter; Trans. Brit. Mycol. Soc., 44: 163-176, figs. I a-s, 2 a-k, Pl. 12.
1961. Saprophytic; Cosmarium and other desmids; plankton of lakes; E.L.D., U.K.

Rhizophydium undatum O.G. Golubeva; Novosti Sistematiki Nizshikh Rastenii, 25: 81-86, fig.
3. 1988. Saprophytic; pine pollen; Lake Teletzkoje; Altiaca Province, Russia

Rhizophydium undulatum Sparrow & Lange; Canad. J. Bot., 55: 1879-1890, figs. 9-11. 1977
Saprophytic; pine pollen bait; Johnson Road Bog; Cheboygan Co., MI, USA

Rhizophydium urcelolatum S.N. Dasgupta & R. John; Bull. Bot. Surv. India, 30: 1-82, fig. 19.
1988. Parasitic; Blastocladia on fruit bait; tank sediment; Lucknow U., India

172

Rhizophydium venezuelensis Karling; Nova Hedwigia, 34: 645-668, figs. 1-9. 1981
Saprophytic; cellophane, snake skin, hemp; soil sample; Venezuela

Rhizophydium venustum Canter; Trans. Brit. Mycol. Soc., 46: 305-320, fig. 7, a-i. 1963
= Zygorhizidium venustum (Canter) Canter 1971
Parasitic; cysts; Uroglena americana; lakes; E.L.D., U.K.

Rozella Cornu 1872; D.J.S. Barr placed provisionally in Olpidiaceae [S]; HP, p. 463. 1990.

- For review of Rozella and Rozellopsis see A. A. Held; Bot. Review, 47: 451-515, 1981.

Rozella blastocladiae in S.N. Dasgupta; Indian Phytopathol. 35: 215. 1982; ? nomen nudum

Rozella coleochaetis Sparrow; Pap. Michigan Acad. Sci., 50: 115-123, figs. A-E, 1964
Parasitic; Coleochaete sp.; Smith's Bog; Cheboygan Co., MI

Rozella diplophlyctidis Karling; Nova Hedwigia, 45: 529-535, figs. 1-6. 1987
Parasitic; Diplophlyctis intestina; moribund Nitella flexilis,; Ross Biological Reserve

Rozella itersoniliae D.J.S. Barr & Bandoni; Mycologia, 71: 1261-1264, figs. 1-4. 1979
Parasitic; Itersonilia perplexante (Hyphomycete); Taraxacum leaf over agar

Rozella longicollis Karling; Sydowia, 19: 213-226, Pl. 46, figs. 21-29. 1965
Parasitic; Pythium (Oomycete); soil sample; N.Z.

Rozella longisporangia Willoughby and Rigg; Nova Hedwigia, 37: 369-383, fig. 1 A-F. 1983
Parasitic; Pythium (Oomycete); field soil with neutral pH; Majorca

Rozella parva Canter; J. Linn. Soc., Bot., 62: 267-278, fig. 3. 1969
Parasitic; Zygorhizidium affluens; lakes; E.L.D., U.K.

RUMINOMYCES Y.W. Ho [N]; Mycotaxon, 38: 397-405. 1990 = Anaeromyces Breton et al.
1990 (described first)

Ruminomyces elegans Y.W. Ho; Mycotaxon, 38: 397-405, figs. 1-3. 1990 = Anaeromyces
elegans (Y.W. Ho) Y.W. Ho 1993. Saprophytic; oat chaff; ramen, Hereford steer; Armindale,
Australia

Scherffeliomyces appendiculatus (Zopf) Sparrow 1936 = Dangeardia appendiculata (Zopf)
Batko; Acta Mycol., 6: 407-435. 1970

Scherffeliomyces leptorrhizus Johns 1956 = Dangeardiana leptorrhiza (Johns) A. Batko; Acta
Mycol., 6: 407-435. 1970

SCHERFFELIOMYCOPSIS Geitler [C]; Oesterr. Bot. Z., 109: 205-275. 1962

Scherffeliomycopsis coleochaetis Geitler; Oesterr. Bot. Z., 109: 250-275, figs. 1-8. 1962
Parasitic ?; Coleochaete soluta & irregula; littoral; Lunzer Untersee, lower Austria

Septochytrium willoughbyi 1.J. Dogma; Nova Hedwigia, 24: 367-377, figs. 1-22. 1973
Saprophytic; lens paper, onion skin & cellophane; soil; virgin forest, Rondonia, Brazil

SEPTOSPERMA Whiffen ex Seymour [C]; Mycologia, 63: 83-93, 1971
= Septosperma Whiffen 1942 (no Latin diagnosis)

For nomenclatural review see Blackwell & Powell; Mycotaxon 42: 43-52. 1991

Septosperma anomalum (Couch) Whiffen ex Seymour (IF 6:291); Mycologia, 63: 83-93, 1971
= Septosperma anomala (Couch) Whiffen 1942 (genus not valid when combination made)

Septosperma irregulare (Karling) I.J. Dogma (IF 4:318); Nova Hedwigia, 25: 121-141. 1974
= Asterophlyctis irregularis Karling 1967

Septosperma multiforme Canter ex Milanez (IF 3:258); Rickia, 6: 63-70. 1974 = Septosperma
multiforma Canter (species described before genus validated); Trans. Brit. Mycol. Soc., 46:
305-320, figs. 4-7. 1963. Parasitic; developing cysts of algae; lakes; E.L.D., U.K.

Septosperma rhizophydii Whiffen ex R.L. Seymour; Mycologia, 63: 83-93, 1971 = Septosperma
rhizophidii Whiffen 1942 (no Latin); Emended by W. Blackwell and M. Powell; Mycotaxon,
42: 43-52. 1991. Also see IF 6:291.

Septosperma spinosum Willoughby ex Milanez; Rickia, 6: 63-70. 1974; = Septosperma spinosa
Willoughby (species described before genus validated); Arch. Mikrobiol., 52: 101-131, fig. 5
a-h. 1965; Parasitic; Rhizophydium coronum; heath; Victoria, Australia

Siphonaria variabilis Petersen 1903 emend. I.J. Dogma; Nova Hedwigia, 25: 107-119. 1974

SOROCHYTRIACEAE Dewel [B]; Canad. J. Bot., 63: 1525-1534. 1985

SOROCHYTRIUM Dewel [B]; Canad. J. Bot., 63: 1525-1534. 1985

7S

Sorochytrium milnesiophthora Dewel; Canad. J. Bot., 63: 1525-1534, figs. 1-36. 1985
Parasitic; tardigrade (Milnesium tardigradum) on moss; bridge & wall; VA, USA
SPARROWIA Willoughby [C]; Nova Hedwigia, 5: 335-340. 1963
Emended by I.J. Dogma; Nova Hedwigia, 19: 503-509. 1970
Sparrowia parasitica Willoughby; Nova Hedwigia, 5: 335-340. Pl. 52. 1963
Parasitic; oogonia of a watermold; soil; north of Thirlmere, E.L.D., U.K.
Sparrowia subcruciformis 1.J. Dogma; Nova Hedwigia, 19: 503-509, figs. 1-16. 1970
Parasitic; various chytrids; forest soil, leaf litter; Cross Village, MI, USA
Sparrowmyces in S.N. Dasgupta; Indian Phytopathol. 35: 206. 1982; ? nomen nudum
Sparrowmyces sparrowii in S.N. Dasgupta; Indian Phytopathol. 35: 206. 1982; ? nomen nudum
Sphaerita dinobryi Canter (IF 3:516); J. Elisha Mitchell Sci. Soc., 84: 56-61, fig. 1, &
Pls. 1, 2. 1968. Parasitic; Dinobryon sertularia; lakes; E.L.D., U.K.
SPIZELLOMYCES D.J.S. Barr [S]; Canad. J. Bot., 58: 2380-2394. 1980
= Phlyctochytrium (in part)
Spizellomyces acuminatus (D.J.S. Barr) D.J.S. Barr; Canad. J. Bot., 62: 1171-1201. 1984
= Phlyctochytrium acuminatum Barr 1969; = P. africanum Gaertner 1954
Spizellomyces dolichospermus D.J.S. Barr; Canad. J. Bot., 62: 1171-1201, figs. 92-100. 1984
Saprophytic; pollen?; soils; widespread
Spizellomyces kniepii Gaertner ex D.J.S. Barr; Canad. J. Bot., 62: 1171-1201. 1984
= Phlyctochytrium kniepii Gaertner 1954 (not valid)
Saprophytic; fly legs, pollen; soil; Republic of South Africa
Spizellomyces lactosolyticus D.J.S. Barr; Canad. J. Bot., 62: 1171-1201, figs. 101-109. 1984
Saprophytic; pollen; soils; Canada
Spizellomyces palustris Gaertner ex D.J.S. Barr; Canad. J. Bot., 62: 1171-1201. 1984
= Phlyctochytrium palustre Gaertner, 1954 (not valid)
Saprophytic; pollen; ditch, Holsburg Moor; Germany; Canada
Spizellomyces plurigibbosus (D.J.S. Barr) D.J.S. Barr; Canad. J. Bot., 62: 1171-1201. 1984
= Phlyctochytrium plurigibbosum D. Barr 1969
Spizellomyces pseudodichotomus D.J.S. Barr; Canad. J. Bot., 62: 1171-1201, figs. 110-120.
1984. Saprophytic; pollen; soils; Canada, U.K., and Mexico
Spizellomyces punctatus (Koch) D.J.S. Barr; Canad. J. Bot., 58: 2380-2394. 1980
= Phlyctochytrium punctatum Koch 1957; = Phlyctochytrium verruculosum Knox 1970;
= P. peruvianum Knox 1970; D.J.S. Barr; Canad. J. Bot., 62: 1171-1201, figs. 110-120. 1984
SPIZELLOMYCETACEAE D.J.S. Barr [S]; Canad. J. Bot., 58: 2380-2394. 1980
See D.J.S. Barr (Canad. J. Bot., 62: 1171-1201. 1984) for key to genera.
SPIZELLOMYCETALES D.J.S. Barr; Canad. J. Bot., 58: 2380-2394. 1980 = Chytridiales
sensu Sparrow (in part). Emended by D.J.S. Barr; Canad. J. Bot., 61: 295-307. 1983
Sporophlyctidium neustonicum Sparrow; Mycologia, 70: 140-149, figs. 15-25. 1978
Parasitic; living pine pollen; neuston; Newnan's Lake, FL, USA
Synchytrium; not included, see Karling, J.S. 1964. Synchytrium. Academic Press, New York
TRIPARTICALCAR D.J.S. Barr [S]; Canad. J. Bot., 58: 2380-2394. 1980
= Phlyctochytrium (in part)
Triparticalcar arcticum (D.J.S. Barr) D.J.S. Barr; Canad. J. Bot., 58: 2380-2394. 1980
= Phlyctochytrium arcticum D.J.S. Barr 1970
Urophlyctaceae Hadar (Compt. Rend. Hebd. Séances Acad. Sci., Ser. 3, 294: 329-333. 1982)
placed into Spizellomycetales (TEM evidence is unclear)
Zygorhizidium affluens Canter; J. Linn. Soc., Bot., 62: 267-278, fig. 2. 1969
= Rhizophydium planktonicum Canter (in part) 1948
Parasitic; Asterionella formosa; plankton of lakes; E.L.D., U.K.
Zygorhizidium asterionellae Pongratz; Schweiz. Z. Hydrol., 28: 104-132, Pl. 3, 19-24. 1966
Parasitic; Asterionella formosa; plankton; Lake Léman, Switzerland
Zygorhizidium chlorophycidis Canter; Nova Hedwigia, 5: 1-6, Pls. 1, 2. 1963
Parasitic; colonial planktonic greens; lakes; E.L.D., U.K.

174

Zygorhizidium cystogenum Canter, Trans. Brit. Mycol. Soc., 46: 305-320, figs. 1, 2. 1963
Parasitic; Dinobryon spp., Uroglena sp.; lakes, E.L.D., U.K.

Zygorhizidium melosirae Canter 1950; Canter emended; J. Linn. Soc., Bot. 60: 85-96. 1967

Zygorhizidium planktonicum forma asterionellae Canter, Jaworski, & Beakes, Nova Hedwigia,
55: 437-455. 1992. Parasitic; Asterionella; lakes; E.L.D., U.K.

Zygorhizidium planktonicum forma synedrae Canter, Nova Hedwigia, 55: 437-455. 1992.
Parasitic; Synedra; lakes, E.L.D., U.K.

Zygorhizidium vaucheriae Rieth; Biol. Zentralbl., 86: 433-448, figs. 1, 2, Pls. 1, 2. 1967
Parasitic; Vaucheria; ditch; Gatersleben, Germany & Tadzhik S.S.R.

Zygorhizidium venustum (Canter) Canter; Nova Hedwigia, 21: 577-592. 1971
= Rhizophydium venustum Canter 1963

MYCOTAXON

Volume LX, pp. 175-180 October-December 1996

COCCOIDELLA EXOCARPI SP. NOV. ON EXOCARPOS SPP.
FROM AUSTRALIA

Z. Q. Yuan’”, C. Mohammed?” and T. J. Wardlaw®

' Dept. of Agric. Science, University of Tasmania, GPO Box 252C Hobart,
Tasmania, Australia, 7001
* CSIRO, Div. of Forestry, Tasmanian Research Centre, Locked Bag No 2, Sandy
Bay Post Office, Sandy Bay, Tasmania, Australia, 7005
3h orestry Commission, 199 Macquarie St. Hobart, Tasmania Australia, 7000

SUMMARY

Coccoidella exocarpt Z.Q. Yuan sp. nov., on living
twigs of Exocarpos spp. is described and illustrated. The
differences between the new species and the known
species of the genus are discussed and the measurements
of the main characteristics of all the known species are
given in a summary table. Differences between
Coccoidella and Microcyclus are briefly discussed.

Exocarpos Labill is a small genus in Santalaceae containing about 20
species distributed across the countries of Australia, Malaysia, Polynesia, New
Zealand and Madagascar. They are almost always parasitic on the roots of other
plants (Curtis 1967). E. cupressiformis Labill, commonly known as the “native
cherry” in Tasmania, is an indigenous species common throughout the more open
eucalypt forests of Tasmania and other temperate Australian regions (Curtis 1967).
Little is known about the fungi which occur on this tree genus. The only fungi
recorded to date on members of this genus are Marasmius exocarpi Berk. and
Meliola exocarpi H. S. Yates, both on E. latifolius R. Br. (Saccardo 1887,
Saccardo & Trotter 1926). In addition, two lichens, Hypogymnia subphysodes
(Krempelh.) Filson var. austerodioides Elix (Brunonia 2, 230, 1979) and
Menegazzia confusa P.W. James (Lichenologist 19, 26, 1987) have been recorded
on the bark of E. cupressiformis from South Australia.

An ascomyceteous fungus appears to be common on E. cupressiformis in
Tasmania. Morphological examination shows that it is a member of the genus
Coccoidella Hohnel. Coccoidella, syn. Coccodothella Theiss. et Syd., was
established by Héhnel (Sitzb. Akad. Wien 118, 847,1909), based on C. scutula

176

(Berk. et Curt.) Héhnel and currently includes eight species (Miller & Arx 1962;
Bose & Miiller 1964; Sivanesan 1987). Species of the genus have superficial disc-
like or crustose stromata, which usually contain a few, fasciculate asci in locules,
and taper to a central hypostromatic foot immersed in host tissue. Ascospores are
ellipsoid or fusoid, hyaline to brown, usually with one median septum, but
sometimes 3-septate. The Australian National Collection of Fungi (NCOF)
database was checked for other fungal collections on Exocarpos. Three collections
referred to as Microcyclus sp., all on E. stictus R.Br. were found in the Herbarium,
Institute for Horticultural Development Victoria (VPRI 17831b, 17832a, 17834a).
Examination of these collections in VPRI show they were identical, particularly as
regards ascospore morphology, to the collections on FE. cupressiformis. As this
fungus has the typical characteristics of the genus Coccoidella mentioned above,
and is not conspecific to any known species, it is proposed here as a new species.
The holotype is deposited in the Herbarium, Department of Primary Industries,
Queensland (BRIP).

Coccoidella exocarpi sp. nov. Z.Q. Yuan Fig. 1, a-d

Stromata caulicola, gregaria, superficialia, cum basibus immersis,
rotunda vel pulvinata, 250-500 um lata, 150-250 um alta, 2-5 (vulge 3) ascomata
continentia. Textum stromatis pseudoparenchymatum, e cellulis atrobrunneis,
crassitunicatis, angularibus ad subsphaericis (5-9 um diam.) et basi cellulis
elongatis compositum. Loculi ascomati sphaerici vel subsphaerici, 100-125 um
lati, 90-100 um alti. Paries loculi 20-40 pm crassus, e cellulis crassitunicatis,
elongatis (7-12 x 2.5-5 pm), in 4-5 stratis disposidis compositus. Asci 90-100 x
18-25 um, bitunicati, obclavati vel ovoidei, ad basim latiores transeuntes, stipitati,
octospori. Pseudoparaphyses absentes. Ascosporae 25-38 x 7-9 um, superimposite
bi- vel multiseriatae, ellipsoideae ad fusoideae cum extremio rotundato, rectae vel
leviter curvatae, 1-3-septatae, non- vel raro constrictae ad septum medium,
hyalinae ad pallide brunneae vel brunneae ad maturitatem.

Holotypus in ramulis (foliis) vividis Exocarpi cupressiformis Labill,
Hobart, Tasmania, Australia, V. 1995, T. Wardlaw, C. Mohammed & Z.Q. Yuan,
BRIP 22785.

Stromata caulicolous, longitudinally aggregated along stems with bases
immersed in the host tissue, usually circular or pulvinate, 250-500 fm wide and
150-250 pm high, containing 2-5 (mostly 3) ascomata. Stromatal tissue composed
of thick-walled dark brown, angular to subspherical pseudoparenchymatous cells
measuring 5-9 tim diam. and elongated at the base of stromata. Ascomatal locules
spherical or subspherical, 100-125 tm wide and 90-100 um high, wall 20-40 um
in thickness, composed of 4-5 layers of thick-walled, elongated cells (7-12 x 2.5-
Sum). Asci 90-100 x 18-25 um (mean = 94 x 22 um, n = 50), bitunicate,
obclavate to ovoid, tending wider toward the base, abruptly short-stipitate,
measuring 7-9 x 3-4 um, 8-spored. Pseudoparaphyses absent. Ascospores 25-38 x

Fig. 1. Coccoidella exocarpi: a. habit, b. ascospores, c. asci, d. stromata in vertical
section. (standard line = 40 um for b and c; 100 pm for d; habit sketch not scale)

177

\
IN

wwe
et
AY

>.
5}
iis
<=

a g

178

7-9 um (mean = 30 x 7.8 pm, n = 50), overlapping bi- to multiseriate, ellipsoid to
fusoid with rounded ends, sometimes slightly narrower toward lower parts, straight
or slightly curved, l-septate, not or slightly constricted at the middle septum,
hyaline to pale brown, guttulate when young, becoming 3-septate and brown at
maturity.

On living twigs (scaled leaves) of Exocarpos spp.

SPECIMENS EXAMINED Australia: Hobart, Tasmania, Exocarpos
cupressiformis, May 1995 T. Wardlaw, C. Mohammed & Z.Q. Yuan (Holotype
BRIP 22785), Wonboyn, New South Wales, E. strictus, 26 March 1992, I. Pascoe
(VPRI 17831b, VPRI 17832a, VPRI 17834a).

Measurements of the main characteristics of all the known species of
Coccoidella are compared in Table 1. Of all the Coccoidella species known
(Miller & Arx 1962; Bose & Miller 1964; Sivanesan 1987), this fungus is closest
to C. perseae A. Sivanesan in ascospore morphology, but differs in having smaller
stromata and much wider asci and ascospores (Table 1). The asci in C. perseae are
cylindric-clavate (Sivanesan 1987), rather than obclavate to ovoid as in this fungus.

Coccoidella is morphologically close to Microcyclus and differs from the
latter genus mainly in the type of stromata and in the number of septa and
pigmentation of the ascospores. Stromata are erumpent from substrate in
Microcyclus and superficial with free hyphae in Coccoidella (Barr, 1987). Based
on early reports (Clements & Shear 1931; Luttrell 1973; Wehmeyer 1975) the
ascospores in both genera are 2-celled, but hyaline in Microcyclus and yellowish to
pale brown or yellow brown in Coccoidella. Sivanesan (1987) described one
Coccoidella species with 1-3-septate, pigmented ascospores. In terms of
pseudoparaphyses, Clements & Shear (1931), Luttrell (1973) and Wehmeyer
(1975) considered that Microcyclus has pseudoparaphyses and Coccoidella lacks
them. Sivanesan (1984) mentioned that Microcyclus has filiform, hyaline
pseudoparaphysoids, usually disappearing in mature stromata. Miller & Sanwal
(1954) noticed remnants of paraphysoid tissue or interthecial tissues in
Microcyclus, and Barr found such remnants in specimens of both M. tinctorius
(Tul.) von Arx and C. scutula, and included both genera in the Dothideaceae (Barr
1987). Based mainly on the characteristics of stroma, ascospore septum and
colour, the present fungus on Exocarpos is best included in the genus Coccoidella.
In addition, the genus Microcyclus is tropical and subtropical (Cannon et al. 1995)
and the ostioles are schizogenous and lined with short periphyses (Barr 1995, pers.
comm.). In the collections of VPRI named Microcyclus sp., we found superficial
stromata and four-celled, pigmented ascospores which are identical to those in our
collections. We therefore assign tho8e collections to the genus Coccoidella along
with our collections in the present study.

cil

‘Apnys Juasord oy}, pur (L861) wesoueats, “(796 1) XIV F JINN, (p96 1) JOIIWIN ® OS0g, Uo paseg,

08S S X67 6l
0°9-0'F X E781
0S-07 X 9I-FI
08S 9X ECOL
SrOPr X LE-0€
GL09X SE-PT
OL-0'S X 0€-8I
09-05 XLI-FI
06-0 L X 8E-ST

0 0CVI X 09-SS
081-91 X OL-0S
OCI-OL X SS-SP
0-07-9I X 09-@r
SEl-Cl X 06-SL

8I-VI X OL-0S

SI-cl X O€T-O0L
Sc-81 X 00-06

OIT-09
00C-0S 1
$C1-06

00S-0S7 X ODOI-0S7
008-009
$L-09 X OST-OFI
00-007 X 0007-00$
O£t-0C7 X OOTI-OLL

0S7C-00C X 000-009
OS7-OSI X 00S-0S7

TWN “a (7g 18 JID) viuosuaagys “_,
UYOH (sng “WW 19 Weg) vjnjnos ‘5,
TIM “A 2 880g “YS vYofiasanb 2;
‘JID 19 “Nog vjoutajnd Dy

‘URATS avasiad *),

‘JID 0 “og vsodsisnf Ds

‘PAS J “SSTOYL (UUaH) 191f “OD,

[eMuUeBS }9 “TINA “A (OW) viddjsna oy
ueny ‘O'Z Idunz0xa ‘D,

(ON).
alodsoose
jo umjdas

(wr)
WIPLM X sus]

olodsoosy

(url)

snosy

“UreIp

g]N00T

(ur)
ISTH X WPI

eUIONSsoosy

x0]]2p10990) JO satdads UMOUY 9} PUL 1dspI0xXa D]Jap10II0FD JO SISTAI}IVACYD UlVU JO JUSWaINsvaA, :] B[qQU]

180

Acknowledgments

The authors are much obliged to Dr. M.E. Barr (Sidney, Canada) for
reviewing the manuscript and to Dr. I. Pascoe of VPRI for loan of materials. The
senior author also wish to thank the University of Tasmania and CSIRO, Division
of Forestry for scholarships.

References

Barr, M.E. 1987. Prodromus to class Loculoascomycetes. Publ. by the author,
Amherst, Massachusetts.

Bose, S.K. and E. Miller. 1964. Central Himalayan fungi - 1. Indian Phytopathol.
17, 3-22.

Cannon, P.F., C.C. Carmaran and A.J. Romero. 1995. Studies on biotrophic fungi
from Argentina: Microcyclus porlieriae, with a key to South American
species of Microcyclus. Mycol. Res. 99, 353-350.

Clements, F.E. and C.L. Shear. 1931. The genera of fungi. Hafner Publ. Co. New
York.

Curtis, W.M. 1967. The student’s flora of Tasmania, Part 3: Angiosperme:
Plumbaginaceae to Salicaceae. Government Printer, Tasmania.

Luttrell, E.S. 1973. Loculoascomycetes. In The Fungi IVA (ed. G.C. Ainsworth,
F.K. Sparrow & A.S. Sussman), pp. 135-219. New York and London:
Academic Press.

Miller, E. and J.A. von Arx. 1962. Die Gattungen der didymosporen
Pyrenomyceten. Beitr. Kryptogamenfl. Schweiz 11, 1-922.

Miller, E. and B.D. Sanwal. 1954. Uber die Gattungen Microcyclus Sacc.,
Coccoidella v. Héhn., Coccodothis Theiss. et Syd. und Coccodothella
Theiss. et Syd.. Sydowia 8, 231-244.

Saccardo, P.A. 1887. Agaricineae, Leucosporae, Marasmius. Sylloge Fungorum. 5,
503-569.

Saccardo, P.A. and A. Trotter. 1926. Pyrenomycetae, Perisporiaceae, Meliola.

Sylloge Fungorum 24, 263-358.

Sivanesan, A. 1984. The bitunicate ascomycetes and their anamorphs. J. Cramer,
Vaduz.

Sivanesan, A. 1987. Coccoidella perseae sp. nov. and its anamorph Colletogloeum
perseae sp. nov.. Trans. Brit. Mycol. Soc. 89, 265-270.

Wehmeyer, L.E. 1975. The Pyrenomycetous fungi. Mycologia Memoir 6, 1-248.

MYCOTAXON

cholo PEGE PREP Ae) a Ta EIN Na IR i EN
Volume LX, pp. 181-190 October-December 1996

NEW AND INTERESTING WOOD-INHABITING FUNGI
(BASIDIOMYCOTINA - APHYLLOPHORALES)
FROM ETHIOPIA

Kurt Hjortstam
Malaregatan 12, S-441 35 Alingsas, Sweden

Leif Ryvarden
Univ. of Oslo, Division of Botany, P. O. Box 1045, N-0316
Oslo, Norway

ABSTRACT

15 species of Corticiaceae s.l. are reported as new to Ethiopia:
Diplomitoporus rimosus (Murrmill) Gilb. & Ryvarden and
Physisporinus rivulosus (Berk. & M.A. Curtis) Ryvarden as
new to Africa. Mycoacia brunneofusca Hjortstam & Ryvarden
and Vuilleminia obducens Hjortstam & Ryvarden are
described as new. Keys are offered to tropical species of
Mycoacia and Vuilleminia.

Keywords: Aphyllophorales, Corticioid fungi, Ethiopia

INTRODUCTION

Ethiopia's wood-inhabiting fungi are poorly known. Bresadola (1896) was
the first to report wood-inhabiting fungi from the country, concentrating
almost exclusively on polypores. Scattered observations were published by
Hennings (1893, 1901, 1904 and 1905). More recently many species of
polypores were reported as new for Ethiopia by Ryvarden & Johansen
(1980), while a number of corticioid species were reported as new from the
country by Hjortstam (1983 and 1987). G. Langer (1994) described
Tofispora repetospora G. Langer & Ryvarden based on an Ethiopian
collection and recently Boidin & Lanquetin (1995) reported 16 corticioid
species as well as describing three new species.

182

In July 1990, one of us (LR) had the opportunity to collect in the central and
southern part of Ethiopia a total of 700 specimens of wood-inhabiting fungi
were obtained. Some agarics and ascomycetes were sent to the Royal Botanic
Gardens, Kew, while some of the more interesting wood-inhabiting
corticioid and poroid species are reported here.

The following localities were visited:

Shoa province, Chilimo, Ginchi, 2400 m, 10 July, Ryv. 28046-28139.

Shoa province, Mena Gesha Nat. Park. 2400 m, 12-14 July, Ryv. 28140-
28308, 28677-28743.

Arussi province, Munessa forest, Lake Langano, 2100m, 15 July, Ryv.
28309-28433.

Arussi province, Wondo Genet, 16-19 July, 2100m, Ryv. 28434-28676.

From the collection number, it is easy to find the exact locality for each
species.

CORTICIOID FUNGI

Athelopsis glaucina (Bres.) Hjortstam

A rather well known fungus on ferns, see Hjortstam (1991) and Hjortstam
and Larsson (1994),

Specimen: Ryv. 28089,

Dextrinodontia molliuscula Hjortstam & Ryvarden
Previously known from Tanzania and Kenya.
Specimens: Ryv. 28296/b, 28342 and 28466.

Gloiothele lamellosa (Henn.) Bres.

This species was originally described from Tanzania and seems to be
restricted to Africa. Specimens have been seen from Malawi and it was also
described as Vesiculomyces epitheloides Boidin et al. (Boidin and
Lanquetin, 1983) from Madagascar. See also Hjortstam (1987) who re-
introduced the generic name Gloiothele Bres.

Specimen: Ryv. No. 28356.

Hyphodontia spathulata (Schrad.:Fr.) Parmasto

Rather well known and possibly a cosmopolitan species, but previously not
reported from Africa (except the Canary Islands, see Rodriques-Armas et al.
(1992). It is known from Europe, North and South America and was
reported from India by Rattan (1977) and recently from Taiwan by E. Lang-

183

er (1994). It seems to be most common in North America and in the central
part of Europe. There is a description with a superb drawing in Eriksson and
Ryvarden (1976). It should be noted that the torulose gloeocystidia are not
always present, or are rather difficult to find.

Specimen: Ryv. No. 28152/C.

Kavinia alboviridis (Morgan) Gilb. & Budington

A widespread species in the Northern hemisphere and additionally known
from Argentina and India (Hjortstam and Larsson, 1994). Recently also
recorded from Brazil (Laessoe No. 52600).

Specimen: on dead deciduous wood Ryv. 28316.

Litschauerella clematitis (Bourdot & Galzin) J. Erikss. & Ryvarden

The micromorphology of this species is not sufficiently well known.

The two specimens from Ethiopia are similar but No. 28211 has warted
spores (KOH) measuring 7-8 x 7.5-9 um while No. 28303 has slightly
smaller, smooth spores (KOH). Litschauerella gladiola (G. Cunn.) Stalpers
& Buchanan (see Stalpers and Buchanan, 1991) is extremely similar except
for its smooth spores and small asterocystidia. The latter elements were,
however, demonstrated by Boidin and Gilles (1986) in a specimen from
Réunion under the name L. abietis (Bourdot & Galzin) Oberw.

Specimens: on Juniperus procera, Ryv. Nos. 28211 and 28303.

Melzericium udicola (Bourdot) Hauerslev

Previously known from Europe, North America, South America (Argentina)
and the Canary Islands.

Specimen: on dead hardwood, Ryv. No. 28334.

Mycoacia brunneofusca Hjortstam & Ryvarden nov. sp.
Species habitu cum Mycoacia fuscoatra , sed differt cystidiis nullis et sporis
ellipsoideis 4-4.5(-4.8) x 2.2-2.5(-2.8)um

Holotypus: Africa, Ethiopia, Shoa Prov. Chilomo Forest, Ginchu 2400 m, 10
July 1990, on dead wood, Ryv. No. 28105 (QO). Isotypus: (K).

Paratypi: Africa, Ethiopia , Shoa Prov. Mena Gesha nat. park 2400-2600m,
12 July 1990, on dead hardwood, Ryv. Nos. 28156 and 28264 (QO).

BASIDIOME resupinate, closely adnate, forming irregular areas 10-15 x 3-5
cm or sometimes larger.

HYMENOPHORE strongly hydnoid and becoming irregularly creviced, at
first light brown, then darkening to more or less blackish brown.

ACULEI densely crowded, conical, smooth, about 1-1.5 mm _ long,
concolorous with the subiculum, or sometimes with a greyish-white tip.

184

SUBICULUM thin to 0.5-1 mm thick, brown to blackish.

HYPHAL SYSTEM monomitic; hyphae mostly agglutinated, hyaline to pale
brown and more or less parallely arranged in the centre of the aculei and
forming a thin trama, apically often strongly encrusted, thin-walled to more
commonly thick-walled, 2.5-3.54m wide. All hyphae with clamp-
connexions.

CYSTIDIA absent.

BASIDIA narrowly clavate, (15-)18-20(-25)um long, with four sterigmata
and a basal clamp-connexion.

SPORES hyaline, ellipsoid, thin-walled, smooth, 4-4.5(-4.8) x 2.2-2.5(-
2.8)um, non-amyloid.

REMARKS. The basidiomes of this species are strongly reminiscent of
Mycoacia fuscoatra (Fr.:Fr.) Donk which, however, has larger, sub-
cylindrical spores measuring 5-6 x 2.5m and distinct subulate cystidia.

KEY TO TROPICAL AND SUBTROPICAL SPECIES OF MYCOACIA

For Mycoacia-like basidiomes without clamp-connexions, see in the first
case Columnodontia and also some species of Phanerochaete and Phlebia.

1. With strongly encrusted cystidia (metuloids). Canary Islands, Thailand,

Australia and New Zealand...................... nothofagi (G. Cunn.) Ryvarden
1. Cystidia otherwise shaped or absent.....................ccccccsssssscccceeeessseeeeeeeees 2
2. GYSIdia PTOSCNE a gewse-atscccaccaliecietaccteach ea cata ae eee ee eee 3
2. Cystidia absent, but in some species with strongly encrusted hyphae in

the apical part-of the:aculeny eye eacsssscsuissereet nicer. Re eee 5
3. Cystidia subulate to acicular, spores 5-6 x 2-2.5 ym. Canary Islands and

Indias isto es er cna ne Rice raat tscc renin mes ate fuscoatra (Fr.:Fr.) Donk
35, Cystidia: subfusiform +. ODtUSC.).505.01.0/0s-sklvcusiocasac sgn aieiy eseeeect on nee 4

4. Cystidia up to 30 pm long, non-septate, spores 5-6 x 2-2.5 ym, Canary
Islands, Morocco, Ethiopia. Mycoacia subceracea (Wakef.) G. Cunn. is
extremely similar or a SYNOMYM...................cccccsesseseeeeeeeees uda (Fr.) Donk

4. Cystidia (or hyphal ends) septate (with clamp-connexions), projecting to
90 pm above the basidia, spores 5-6 x 2.5-3 um (Cunningham 1959).

New Zealand seers. cos cutscstaiicss Mer tare lutea (G. Cunn.) Hjortstam
5. Aculei with strongly encrusted and projecting hyphal ends..................... 6
5. Encrusted hyphae not prominent or absent......................cccceseeeeececeeeeeeees 8

6. Spores ellipsoid, 4.5-6 x 2.5-3 um. New Zealand.
5 dil PERERA RCE sees, BANE sg) soe cere Lg tered columellifera (G. Cunn.) Jiilich

185

7. Spores 4-4.5 x 1.8-2 pm. Australia, New Zealand (Ecuador?)

Te A UPA EEA 11h DALLES ODT Ri LH subfascicularia (Wakef.) Hjortstam
8. Spores suballantoid to allantoid............ eee ee eeneecesseeeceesneeeeeenaeees 9
BP SPOLes CLIIDSOL LH Hic .c ot cite ieces Oe OMA MO LaaNae Nia bah vessTabdone Mandaidee te 10
9. Spores allantoid, 4-4.5 x 1-1.25 pm. Brazil

9. Spores (sub)allantoid, 3.5-5.5 x 1.5-2 ym. Canary Islands, Africa and

10. Basidiome light brown to blackish brown, teeth conical, spores 4-4.8 x
2.2-2.8 pm. Ethiopia....................... brunneofusca Hjortstam & Ryvarden

10. Basidiome pale buff to cream, teeth cylindrical, spores 4-4.5 x 2-2.5 um
(Gilbertson et al. 1976). USA (Arizona)........... austro-occidentalis Canf.

Phanerochaete rosea (Henn.) P. K. Buchanan & Hood

This is a well known species, at least in eastern Africa, and has been
previously reported by Hjortstam and Ryvarden (1980) from Tanzania and
Kenya. It also seems to be distributed in the southern part of Africa and has
been recently reported from New Zealand by Buchanan and Hood (1992).
The micromorphology conforms with the generic concept of Phanerochaete,
above all by the absence of clamp-connexions and the morphology of the
basidia and hyphae. For a modern description with figure see Hjortstam and
Ryvarden (1980).

Specimen: Ethiopia, Shoa Prov. Wondo Genet Forestry College, near
Shashemene 1800 m, 18 July 1990, on dead hardwood, Ryv. No. 28487.

Phlebiella ardosiaca (Bourdot & Galzin) K.H. Larss. & Hjortstam

A little known fungus, mainly distributed in central and southern Europe
The species is further known from the Canary Islands (Rodriques-Armas et
al. 1992) and has been collected by Ryvarden from Malawi.

Specimen: Ethiopia, Shoa Prov. Mena Gesha nat. park. 2400-2600 m , 12
July 1990, on Juniperus procera, Ryv. No. 28187.

Repetobasidiellum fusisporum J.Erikss.& Hjortstam

Previously known from Sweden and Brazil, this is an obligate fungus on
different kinds of ferns (in South America collected on Cyathea sp. and
Dicksonia sp.). It grows on remnants still attached to the plant and is often
found together with Athelopsis lembospora (Bourdot) Oberw. The material
from Ethiopia is poorly developed, or rather too dry, but some typical spores
and a few dendrohyphidia have been observed. As the fungus was
insufficiently known when described, we offer a more detailed description
based on specimens from Brazil. A good drawing was made by John
Eriksson (in Eriksson, Hjortstam and Ryvarden, 1981), and it should also be

186

noted that the species was neotypified by Hjortstam (1987).

BASIDIOME resupinate, closely adnate, thin to moderately thick, at first
pruinose and discontinuous, then continuous and ceraceous, whitish to more
commonly grey and especially when old more or less glossy.

HYPHAL SYSTEM monomitic; hyphae with clamp-connexions, 1.5-2.5(-
3)um wide, often strongly agglutinated and forming a very dense tissue. In
some specimens clamp-connexions are difficult to discern without making a
thin, vertical section of the fungus.

CYSTIDIA absent.

DENDROHYPHIDIA present in most cases and easily observed, distinctly
ramified and protruding as much as 10um above the basidia, thin-walled,
widened in the basal part and often strongly encrusted.

BASIDIA at first subglobose, then subcylindrical to slightly urniform,
mostly constricted in the middle part, 20-30 x (6-)7-8um, with four
sterigmata and a basal clamp-connexion, the repetition (mentioned in the
original description) as well as clamp-connexions are difficult to observe
except in fresh material in either Melzer’s reagent or Cotton blue.

SPORES mostly sublunate to subfusiform, (6.5-)8-12 x 2.5-3.5m (in the
original description 6.5-8 x 2.5-3um), thin-walled, smooth, non-amyloid,
non-dextrinoid and cyanophilous.

Specimen: on Adianthum sp. Ryv. No. 28468.

Scytinostromella heterogenea (Bourdot & Galzin) Parmasto

This species is mainly distributed in the Northern Hemisphere (Eriksson,
Hjortstam and Ryvarden, 1984) and is also known from Morocco
(Malencon, 1982) and India (Rattan, 1977).

Specimen: Ryv. No. 28296.

Sistotrema subtrigonospermum D.P. Rogers.
Specimen: Ryv. No. 28048.

Vuilleminia obducens Hjortstam & Ryvarden nov. sp.
Differt a Vuilleminia comedens sporis subglobosis, 9-12(-14) x 7-8(-9) tm.

Holotypus: Africa, Ethiopia, Shoa Prov. Chilomo Forest, Ginchu 2400 m, 10
July 1990, Ryv. No. 28131 (QO). Isotype: (GB, K). Paratype: Ryv. No. 10568

(kK).

BASIDIOME resupinate, effused, 0.2-0.3 mm thick, closely adnate but
loosening at the margin.

HYMENOPHORE smooth to slightly tuberculate, pale brown with a
somewhat violet tint, margin indeterminable or more commonly darker than
the fertile part.

187

HYPHAL SYSTEM monomitic; all hyphae with clamp-connexions,
generative hyphae thin-walled, other hyphae thin-to moderately thick-
walled, hyaline, often encrusted, 3.5-44m wide, basal hyphae agglutinated
and usually very pale yellowish, otherwise similar and forming a thin, 0.1-
0.2 mm deep, pale brown and rather hard subiculum.

DENDROHYPHIDIA indistinctly developed, encrusted.

CYSTIDIA few, tubular and often apically subcapitate, thin-walled or
apically slightly thick-walled, 60-80 (-100)ym.

BASIDIA tubular-clavate, somewhat sinuous, 60-70(-80)um long and about
10,.m wide in the upper part, with four, rather stout sterigmata and a basal
clamp-connexion.

SPORES subglobose, smooth, thin-walled, 10-12(-14) x 7-8(-9)um,
non-amyloid, non-dextrinoid and cyanophilous.

REMARKS. In outward appearance very similar to some species of
Peniophora, e.g. P. quercina (Pers.:Fr.) Cooke, but immediately separated
from this genus under the microscope. It possibly has the same ability as
Vuilleminia comedens (Nees:Fr) Maire to break through the bark or at least
loosen characteristically at the margin. The species should be easily
recognised by its large basidia and by being the only species in Vuilleminia
so far with ellipsoid spores.

KEY TO TROPICAL AND SUBTROPICAL
SPECIES OF VUILLEMINIA

PC YSUIGIA DICSCNE ee Tete cde tthe eect yiak Sue OEE, Mure AE ea 2
Pee @ystidia ADSENEL FF Fee ee Me Se POR PE IT CEE A SY 4
2. Cystidia subulate, spores allantoid, 15-20 x 4.5-6 ym. Canary Islands.
i Siallte hs USUI Let 8. a eas g CAs e cystidiata Parm.
Da CYSUIGIA CADITAleNy sth. niclt te ittee Herta eatin is ea, tate amar Maren 3
3. Cystidia few, tubular, apically subcapitate, spores globose 10-14 x 7-9
LMS thiopias en Geese nts obducens Hjortstam & Ryvarden

3. Cystidia distinctly capitate, spores allantoid 15-20 x 5-6 ym. Morocco.
sth RSAC As RUA StL RR a lees a AALS macrospora (Bres.) Hjortstam

42sespores: clobose to SubglODOSE: 2 Pee ee uO ca sbeh se 5
4yespores allantoid or ellipsoid (5,305.04. aes 6
5. Spores 14-18 x 9-13 ym. Argentina.
VS id te ec a AEE RO Melle en subglobispora Hallenb. & Hjortstam
DOMES POLES LOMLA KP) OPIN, «css sacelecssnnd icetconge sumer eh entice tot ashore obducens
6. Spores allantoid 15-25 x 5-6 ym. Ecuador ?, Morocco, Tunisia, New
CALAN CGT eee hes Le Aa eM ee comedens (Nees:Fr.) Maire

6. Spores ellipsoid 20-28 x 9.5-13 um (Boidin et al. 1994). Cameroon ?
Lg Ae Brae OL Do ok Le Se Mee pea ee a aL Gee erase GALE cai REG SA megalospora Bres.

188

Xenasma pruinosum (Pat.) Donk

A well known though rare species, though rare, from the Northern
Hemisphere, originally described from Tunisia. For further information on
its tropical distribution see Hjortstam and Larsson (1994).

Specimens: Ryv. Nos. 28126 and 28341.

Xenasma pulverulentum (Litsch.) Donk

Distributed mainly in the Northern Hemisphere and originally described
from Austria. Also known from Venezuela (Liberta and Navas, 1978),
Colombia and Argentina (Gomez, 1973) and Reunion (Boidin and Gilles,
1989) and reported from Morocco by Malencon (1982).

Specimen: on Juniperus procera, Ryv. No. 28297.

Xenasma rimicolum (Karst.) Donk.
Also reported by Boidin & Lanquetin (1995).
Specimen: Ryv. 28068.

POROID FUNGI

Diplomitoporus rimosus (Murrill) Gilb. & Ryvarden

New to Africa. The species is only known from Juniperus, as indicated in
Gilbertson & Ryvarden (1986), where a detailed description is given. The
distinct characters are the cracking of the pore surface, hence the name, the
tubular cystidia and the host. It adds to the group of species restricted to
Juniperus such as Pyrofomes demidoffii and Antrodia juniperina both
known from this host in Africa (including Ethiopia), Europe and America.
D. rimosus has not as yet been found in Europe, but it is very

probable that it will be found, taking the distribution of the two other species
into account.

Specimen: on Juniperus procera, Ryv. 28227.

Physisporinus rivulosus (Berk. & M.A. Curtis) Ryvarden
New to Africa. The species is widespread in America and rare in Europe.
Specimen: on dead hardwood, Ryv. 28444,

LITERATURE CITED

Boidin, J. and Gilles, G. 1986. Basidiomycetes Aphyllophorales de I'ile de
la Réunion. II. Les genres Tubulicrinis, Tubulicium et Litschauerella. Bull.
Soc. Mycol. Fr. 102:279-290.

—1989, Basidiomycetes Aphyllophorales de l'ile de la Réunion. XIII et XIV.
Bull. Soc. Mycol. Fr. 105:130-150.

189

Boidin, J. and Lanquetin, P. 1983. Basidiomycetes Aphyllophorales
Epitheloides Etales. Mycotaxon 16:461-499.

—1995. Sur quelques Corticies (Basidiomycotina) de lEthiopie. Crypt.
Mykol. 16:79-84.

Boidin, J., Lanquetin, P. and Gilles, G. 1994. Contribution a la connais-
sance du genre Vuilleminia (Basidiomycotina) Bull. Soc. Mycol. Fr. 110:91-
107.

Bresadola, G. 1896. Contribuzioni alla conoscenza della flora dell‘ Africa
Orientale. Ann. Royal. Inst. Bot. It. Roma 6:177-180.

Buchanan, P.K. and Hood, LA. 1992. New species and new records of
Aphyllophorales (Basidiomycetes) from New Zealand. N. Z. Jl Bot. 30:95-
112.

Cunningham, G.H. 1959, Hydnaceae of New Zealand. Part. II-The Genus
Odontia. Trans. Roy. Soc. New Zealand 86:65-103

Eriksson, J., Hjortstam, K. and Ryvarden, L. 1981. The Corticiaceae of
North Europe Vol.6. Phlebia-Sarcodontia. Fungiflora, Oslo, Norway.
—1984. The Corticiaceae of North Europe Vol. 7. Schizopora - Suillo-
sporium. Fungiflora, Oslo, Norway.

Eriksson, J. and Ryvarden, L. 1976. The Corticiaceae of North Europe.
Vol. 4. Hyphodermella - Mycoacia. Fungiflora, Oslo, Norway.

Gilbertson, R.L., Burdsall, H.H. and Canfield, E.R. 1976. Fungi that de-
cay mesquite in southern Arizona. Mycotaxon 3:487-551.

Gilbertson, R. L. and Ryvarden, L.1986. North American Polypores.
Abortiporus - Lindtneria. Fungiflora, Oslo, Norway.

Goméz, C.E. 1973. Xenasma y generos afinis de los Alrededores de Buenos
Aires (Aphyllophorales). Bol.Soc. Arg.Bot.14:269-281.

Hennings, P. 1893. Fungi Aethiopici Arabici. Bull. herb. Boissier 1:97-122.
—1901. Fungi Africae orientalis I. Engler Bot. Jahrb. 28:318-329.

—1904. Id. III. Engler Bot. Jahrb. 34:39-57.

—190S5. Id. IV Engler Bot. Jahrb. 38:102-118.

Hjortstam, K. 1983. Studies in tropical Corticiaceae (Basidiomycetes) V.
Specimens from East Africa collected by L. Ryvarden. Mycotaxon 17:555-
Dis

—1987. Studies in Tropical Corticiaceae (Basidiomycetes) VII. Specimens
from East Africa, collected by L. Ryvarden II. Mycotaxon 28:19-37.

—1987. A _ check-list to genera and species of corticioid fungi
(Hymenomycetes). Windahlia 17:55-85.

—1991. Athelopsis instead of Pteridomyces (Corticiaceae, Basidiomycetes).
Mycotaxon 42:149-154.

Hjortstam, K. and Larsson, K.H. 1994, Annotated check-list to genera and
species of corticioid fungi (Aphyllophorales, Basidiomycotina) with special
regards to tropical and subtropical areas. Windahlia 21:1-75.

190

Hjortstam, K. and Ryvarden, L. 1980. Studies in Tropical Corticiaceae
(Basidiomycetes) I. Mycotaxon 10:269-287.

Langer, E. 1994, Die Gattung Hyphodontia John Eriksson. Bibl. Mycol.
154:1-298.

Langer, G. 1994. Die Gattung Botryobasidium Donk (Corticiaceae, Basidio
mycetes). Bibl. Mycol. 158:1-457.

Liberta, A. E. and Navas, A. J. 1978. Notes | on Venezuelan Corticiaceae
(Basidiomycetes). Can. Jl Bot. 56:1777-1781.

Malencon, G. 1982. Nouvelles contributions a la flore Se ee du
Maroc-III. Bull. Soc. Mycol. Fr. 98:18-248.

Rattan, S.S. 1977. The resupinate Aphyllophorales of the North Western
Himalayas. Bibl. Mycol. 60:1-427.

Rodriguez-Armas, J.L., Ryvarden, L., Hallenberg, N. and Tejera, E.B.
1992. New and noteworthy species of Aphyllophorales (Basidiomycotina)
from the Canary Islands. Mycotaxon 45:433-447.

Ryvarden, L. and Johansen, I. 1980. A preliminary polypore flora of East
Africa. Fungiflora, Oslo, Norway. 636 pp.

Stalpers, J. A. and Buchanan, P.K. 1991. Type studies of the species of
Pellicularia and Peniophora described by G.H. Cunningham. N.Z. Jl Bot.
29:331-340.

MYCOTAXON

ill anand Sa a II IN a ER a ET Cad! A ie Ss
Volume LX, pp. 191-200 October-December 1996

A NEW SPECIES OF BARTALINIA TASSI
(DEUTEROMYCOTINA, COELOMYCETES)

Anderson, Freda! & Bianchinotti, Virginia
1 Departamento de Agronomia

2Departamento de Biologia y Bioquimica
Universidad Nacional del Sur
San Juan 670 (8000) Bahia Blanca. ARGENTINA.

ABSTRACT

A fungus belonging to the genus Bartalinia Tassi was isolated
from chlorotic leaves of Mimosa rocae (Leguminosae). As it could not
be assigned to any of the known species it is here described as a new
one under the name Bartalinia mellea. This is the first report of the
genus for Argentina.

RESUMEN

Se aislo un hongo perteneciente al genero Bartalinia Tassi a
partir de hojas cloroticas de Mimosa rocae (Leguminosae). Como no
pudo ser asignado a ninguna de las especies conocidas se lo describe
como nueva bajo el nombre Bartalinia mellea. Esta constituye la
primera cita del género para la Republica Argentina.

192

While carrying out research on diseases affecting endemic wild
plant species in the south of Buenos Aires Province (Argentina), a
specimen of Mimosa rocae Lorentz & Niederlain (Leguminosae) was
collected because of the chlorosis observed on some of its leaves. M.
rocae is the southernmost species of the genus and grows only on the
hills in the south of Buenos Aires Province in Argentina and in

Uruguay.

From this material a fungus belonging to the genus Bartalinia
Tassi was isolated, but after close examination it could not be properly
assigned to any of the known species of the genus.

In this paper we provide a full description of Bartalinia mellea
which we propose as a new species.

MATERIALS AND METHODS

Isolation: The fungus was isolated from chlorotic leaves of Mimosa
rocae on Potato Dextrose Agar (PDA). These were superficially
desinfected with a tenfold dilution of a commercial bleacher (NaOCl).

Studies on host tissue: Leaves of M. rocae were treated at a pressure of
one atmosphere for 20 minutes and inoculated later with the fungus.
They were then incubated in a humid chamber at 25°C till fruiting
bodies were formed.

Cultural studies: They were performed both on PDA and Malt Extract
Agar (MA), both prepared according to Dhingra & Sinclair (1985).

Observations, photographs, and measurements of conidiogenous cells
and conidia were made in tap water. Otherwise they were mounted in
KOH and stained with phloxine 5%.

193

RESULTS
Bartalinia mellea Anderson & Bianchinotti, sp. nov.

Etymology: from Latin melleus (honey-coloured) referring to the colour of
conidia.

Conidiomata eustromatica, superficialia, separata_ vel
‘confluentia, forma irregularia, multilocularia, atro brunnea vel
nigra; parietes crassi ex textura angularis, extra ex cellulis brunneis
et intra ex cellulis pallidioribus compositi, parietes interloculares ex
textura porrecta compositi, 76-560 x 76-305 um.

Conidiophora simplicia vel irregulariter ramosa, pauci
septata, hyalina, laevia, ex cellulis interioribus loculorum oriunda, 6-
11 x 2-3 um.

Cellulae conidiogenae holoblasticae, indeterminatae, discretae
vel integratae, ampulliformes, cylindricae, subcylindricae vel
irregulares, 3-13 x (1.5-) 2-3 (-4) um.

Conidia ellyptico-fusiformia, transversa 4-euseptata, ad septa
raro constricta, laevia, (16-) 19-24 (-27.5) x 3-4 (-5) um. In culture
(PDA) (19-) 24-29 (-32) x 3-5 um. Cellula apicalis conica, hyalina,
tenuitunicata, cum appendice singulari inserta, cum 3 rare 2-4
divergentibus, hyalinis, flexibilibus ramis, 14-24 um long. Cellulae
medianae cylindraceae, pallide brunneae vel ochraceae. Cellula
basalis obconica, base truncata, subhyalina, cum appendice singulari
simplici, excentrica, 6-15 um long.

Coloniae in PDA atrovirentes, reverso atro, in MA fumosae,
reverso fusco.

Typus: BBB 158.

Conidiomata stromatic, superficial, separate or confluent,
plunilocular, irregular in shape, dark brown to black, 76-560 x 76-305
uum. Walls thick, composed of bright brown, thick-walled cells disposed
in fextura angularis, paler and thinner in the inner layers. Interlocular
walls composed of thin-walled fextura porrecta. Ostiole absent,
dehiscence irregular by breakdown of the upper wall.

194

Conidiophores lining the inner layer of the conidiomata walls
from which they originate, simple or irregularly branched, septate near
the base, cylindrical to ampulliform, colourless, 6-11 x 2-3 ym.

Conidiogenous cells holoblastic, determinate or indeterminate,
discrete or integrated, cylindrical, subcylindrical, ampulliform or
irregular in shape, colourless, 3-13 x (1.5-) 2-3 (-4) um.

Conidia cylindrical or fusiform, smooth-walled, straight to
slightly curved, 4-euseptate, not (or only very slightly) constricted at
the septa, (16-) 19-24 (-27.5) x 3-4 (-5) um [mean = 21.7 x 3.4 pm].
Lenght/width mean: 6.5. In culture (PDA) (19-) 24-29 (-32) x 3-5 pm.
[mean = 25.2 x 3.7 um; lenght/width mean: 7]

Apical cell hyaline, conic, thin-walled, bearing a 3- (rarely 2 or
4) furcated hyaline appendage drawing out laterally from the apex.
Appendage branches divergent, flexible, tapering gradually towards the
apex, 14-24 um long. [mean = 20 pm]

Median cells cylindrical, ochraceous to pale brown; the one next
to the basal cell is always the longest: lenght/width ratio= 2,6 versus
1,5 for the others.

Basal cell obconic, attenuated at the base, subhyaline, with an
exogenous, laterally inserted, hyaline appendage, 6-15 um long. [mean
= 10 pm]

Colonies in PDA deep green, black on the reverse, growing 4
cm/week. Colonies in MA smoky, brown on the reverse, growing
slightly slower than on PDA. Mycelium immersed, branched, septate.

Type: Conidiomata on leaves inoculated with the fungus isolated from chlorotic
leaves of Mimosa rocae collected from Co. Tres Picos, Villa Ventana, Buenos Aires
Province, Argentina, Nov. 1988, Delhey & Anderson and dried cultures (PDA and
MA) of an isolate from the same material (BBB 158).

Teleomorph: unknown.
Known distribution: Argentina.

Figs. 1-8. Bartalinia mellea. Fig. 1. Vertical section through plurilocular
conidiomata. Fig. 2. Section through interlocular walls. Figs. 3-5.
Conidiophores. Figs. 6-8. Conidia. Notice an apical 4-furcated appendage
(arrow). (Fig. 8. Phase contrast).

Scale bars: Figs. 1-2 = 50 um, Figs. 3-8 = 10 um.

195

196

All the conidial cells have the capacity of germinating, with the
exception of the apical one. This cell is empty in mature conidia.

A comparison between the distribution of the lenghts of conidia
in culture and on inoculated leaves was made (Fig. 10) on the basis of
80 measurements from each substratum. Conidia on PDA were longer,
with higher frequencies between 24-27 um while on leaves the highest
frequencies observed were between 19-24 pm.

Fig. 9. Bartalinia mellea. Fertile hymenium. Scale bar: 10 pm.

197

30
we Culture
25
wy Leaves
20

Frequency
a

mE! iets

16.0 17.6 19.2 20.8 22.4 24.0 25.6 27.2 28.8 30.4 32.0
Conidium lengths (um)

Fig. 10. Comparison between conidium lenghts of Bartalinia mellea
from inoculated leaves of Mimosa rocae and from culture (PDA). B.
robillardoides extreme measurements are indicated with ¥.

DISCUSSION

Among the species included by Nag Raj (1993) in the most
recent review of the genus Bartalinia Tassi, Bartalinia mellea is
closest to Bartalinia robillardoides Tassi for having conidia which are
regularly 4-septate, but differs from it in a number of other
characteristics (Table 1), the most salient being the nature of the
conidiomata and conidiophores. B. robillardoides possesses smaller,
unilocular fruiting bodies and conidiophores which are reduced to
conidiogenous cells.

The production of plurilocular conidiomata, is a feature which B.
mellea shares with B. tamarindi Nag Raj and B. bischofiae Nag Raj,
however these two species have 3-septate conidia.

198

/ |B robillardoides

Conidiomata
to indeterminate
ocule configuration
slobose to angular
dimensions 76-560 x 76-305 jum

Conidiophores
reduced to developed, simple or
conidiogenous cells branched

POC discrae ee | integrated or discrete
shape ampulliform cylindrical to

Conidia

septation 4-septate

wm | =
27 ©
| &
© Oo

Conidiogenous cells

=
=
=
&
S
wn

shape subcylindrical cylindrical or fusiform

colour ochraceous to pale
brown
dimensions (16-)19-24(-27) x
3-3.5 um 3-4(-5) um

mean length/width |

Apical appendage
ranching

oy
N

3-branched
15-18 um

(2-)3(-4)-branched
14-24 wm

— | o>
(TQ

Basal appendage

Table 1: Comparison between Bartalinia robillardoides (fide Nag, Raj, 1993)
and Bartalinia mellea.

199

There are other species in the genus which can produce well
developed conidiophores (viz. B. bischofiae, B. lateripes (Ellis &
Everheart) Nag Raj and B. pistacina (Maas) Nag Raj) but shape, size
and septation of conidia readily distinguish B. mellea from them.

According to our observations conidium lenghts in culture varied
substantially when compared to those on leaves (Fig. 10), but other
features such as conidiomata characteristics, nature of conidiophores,
morphology of conidiogenous cells and conidia remained the same. The
stability of these features strenghtens our trust in their reliability as
basis for the description of this new species.

This is, in our knowledge, the first report of a species of the
genus Bartalinia Tassi in Argentina. B. cordylines (Speg.) Hohnel
cited by Guba (1961) for Argentina is a mistake according to Nag Raj
(1993) since Hohnel never published this combination. The fungus
originally described by Spegazzini (1910) as Pestalozzina cordylines is
today Zetiasplozna cordylines (Speg.) Nag Raj.

The genus has already been recorded in Brazil, but the species
described from that country have not been formally accepted (Nag Raj,
1993).

ACKNOWLEDGMENTS

We are very grateful to Dr. T.R. Nag Raj for having examined
our material and for his very helpful suggestions during the
development of this study. We are also greatly indebted to Lic. Marta
Garelli for improving the Latin diagnosis and to Dr. GH. Boerema for
reviewing our manuscript. We wish to thank Dr. Rolf Delhey very
specially for his constant advice and encouragement .

Partial financial suppport was provided by the CONICET (Consejo
Nacional de Investigaciones Cientificas y Técnicas de la Republica
Argentina) and by the Universidad Nacional del Sur. MVB has a
Posdoctoral Fellowship from the CONICET.

200

LITERATURE CITED

Dhingra, O.D. & Sinclair, J.B. 1985. Basie Plant Pathology Methods.
Boca Raton. Florida. 355 pp.

Guba, E.F. 1961. Monograph of Monochaetia and Pestalotia.
Harvard University Press, Cambridge, Massachusetts, U.S.A.,
342 pp.

Nag Raj, T. R. 1993. Coelomycetous anamorphs with appendage-
bearing conidia. Mycologue Publications, Waterloo, Ontario,
Canada, 1101 pp.

Spegazzini, C. 1910. Mycetes Argentinenses. An. Mus. Nac. Bs. As. 20
(ser. 3 v. 13): 372.

MYCOTAXON

Volume LX, pp. 201-212 October-December 1996

PINATUBO ORYZAE GEN. ET SP. NOV. AND ITS IDENTITY
DURING ROUTINE TESTS OF RICE SEED

J. B. Manandhar! and T. W. Mew2

1Molecular Genetics Research, 21/241 Dilli Bazar, Kathmandu, Nepal and
2international Rice Research Institute, P. O. Box 933, 1099 Manila, Philippines.

Abstract: Pinatubo Manandhar & Mew and P. oryzae Manandhar & Mew are
new genus and species respectively for our new fungus isolated from rice
seed lots grown in the Philippines. P. oryzae produces 2 kinds of hyaline,
oboviform to pyriform sympodio-conidia; 1-septate, dry, 6.7-10.2 x 2.2-3.3
um (average 8.2 x 2.5 um) conidia from solitary conidiophores and O-septate,
wet, 5.6-8.4 x 1.8-2.6 um (average 6.7 x 2.1 tm) conidia from sporodochial
conidiophores on the same niche. In sporodochium, first conidium initiates
on minute denticle near the tip of a conidiophore, second conidium initiates
similarly by elongating the axis just below the first conidial attachment
displacing it to a side, and so on forming a heap of conidia. The fungus is
weakly parasitic to germinating rice seedlings showing chlorotic gray lesions
on plumules. P. oryzae occurs on most seed lots of rice grown in the
Philippines. Higher percentage of the fungus was detected in seed lots
produced during wet than dry season.

Introduction: Our new fungus is frequently encountered in rice seed lots
grown in the Philippines that forms morphologically different sympodio-
conidial anamorphs, solitary conidiophores and sporodochial conidiomata on
the same niche, which we believe it belongs to new genus. Genus Pinatubo
is erected to place this fungus. Arthrobotrys was created by Corda (1839)
for hyphomycetes with uniseptate, hyaline conidia with an apiculiform base,
that develop on sterigmata arranged in whorls on the tips and nodes of
septate conidiophores. The genus was based on the type specimen, A.
superba Corda. Nematode-capturing hyphal networks in A. superba were
described later by Zopf (1888). Cooke and Godfrey (1964) prepared a key
for 12 species of Arthrobotrys having adhesive hyphal branches, networks,
constricting or nonconstricting rings and aerial sticky knobs to capture live
nematodes. Haard (1968) updated the key for 12 species of Arthrobotrys
with uniseptate conidia either borne on nodules or not and with predaceous
organs to capture live nematodes. The synnematous saprobic A. stilbacea
Meyer was excluded by Haard (1968) on the grounds that it probably
belonged to another genus. Barron and Davidson (1972) described A.
anomala Barron & Davidson in Arthrobotrys although it has nonseptate
conidia on the grounds that conidia become uniseptate before germination

202

and the species has adhesive hyphal branches. Jarowaja (1970) and Schenck
et al. (1977) re-evaluated and broadened the generic concept of
Arthrobotrys and amalgamated 31 and 47 species respectively of
nematode-trapping hyphomycetes with distinctly denticulate conidiophores.
Jarowaja (1970) included typically 1-septate hyaline conidia, while Schenck
et al. (1977) included species with O to pleuriseptate hyaline conidia such as
new species A. amerospora Schenck, Kendrick & Pramer. Arthrobotrys
stilbacea is a saprophyte isolated from palm leaves in the former Belgium
Congo (Meyer, 1958), but Jarowaja (1970) and Schenck et al. (1977)
included it within their expanded generic concept of Arthrobotrys without
proof of its nematophagous properties. Oorschot (1985) re-evaluated and
accepted 31 species that includes 5 varieties of Arthrobotrys restricting 0
to 3 septate hyaline ovoid to clavate conidia borne on short denticles.
Synnematous fungi, A. stilbacea_ is excluded but A. dendroides
Kuthubutheen, Muid & Webster included based on the nematode-trapping
nature in the current generic concept of Oorschot (1985). Current
literature review shows A. constringens Dowsett, Reid & Kalkat (Dowsett et
al., 1984), A. ferox Onofri & Tosi (Onofri & Tosi, 1992) and A. pauca
McCulloch (McCulloch, 1977) have been described but have not added to
the expanded genera of Arthrobotrys.

A fungus was frequently encountered in 2-80% of seeds in rice seed lots
grown in the Philippines. This fungus occurs both on germinated and non-
germinated seeds concealing more pathogenic fungi. This fungus appears to
be a new genus Pinatubo and new species P. oryzae and is described
below.

Materials and methods: Fungal structures were picked up directly from
culture plates or from infected or infested grains into a drop of lactophenol
with 0.3% cotton blue or a 1 M KCI on a slide; a cover slip was applied and
sealed with nail polish. Chlamydospores were directly transferred to a drop
of glycerin to examine the color. Fungal structures were measured and
camera lucida drawings were made using a 100x objective of a compound
microscope, except for some solitary conidiophores and pionnotes which
were observed with a 10x objective. The test fungus was grown twice on 5
plates each of potato dextrose agar (PDA) or 2% water agar (WA) at 28°C
under a 12-hr photoperiod and measured its colony diameter in 7 days after
inoculation.

Four-hundred seeds of each of 20 cultivars (IR 8, IR 20, IR 32, IR 36, IR 38,
IR 42, IR 44, IR 48, IR 50, IR 52, IR 54, IR 56, IR 58, IR 62, IR 68, IR 70, IR 72, IR
74, PSBRc 2, and PSBRc 4) grown at the International Rice Research Institute ©
(IRRI) at Los Bafhos, Laguna during 1992 wet season or 1993 dry season
were plated by the wet blotter paper method (Mia et al. 1985; Nath et al.
1970), incubated under continuous near ultraviolet light for 7 days at 25°C,
and examined with a binocular microscope to compare the percent and
degree of infestation due to P. oryzae in dry versus wet season. Degree of
infestation was based on the percentage cover of fungal hyphae on the rice
seeds or seedlings in 7 days, where 0 = none to 9 = 100% surface covered

203

by the fungus. There were 25 seeds per plate, 4 plates per replication and
4 replications.

Individuals of Aphelenchoides besseyi Christie were isolated using the
Baerman funnel method (Southey, 1986) from the rice seeds of line IR
19319-5-3-3-2-1 grown in the 1992 wet season. Thirty nematodes were
surface-sterilized in 1% streptomycin sulfate for 2-3 min and washed 3 times
in sterilized distilled water and then transferred to 3-day old colony of
Alternaria alternata (Fr.) Keissler (isolated from rice seed) grown on PDA as
described by Southey (1986) and incubated at 28°C and a 12-hr
photoperiod for 10 days. Nematodes from A. alternata cultures were then
picked up and surface-sterilized as described above. Thirty nemas per 9-cm
diam plate were placed on a 3-day-old colonies of P. oryzae grown on PDA
or WA to study whether or not this fungus is nematophagous. A small
chunk of an P. oryzae. Colony was cut, transferred to a plate containing
sterile water and approximately 100 nematodes were then picked up
randomly and transferred to a drop of glycerin, a briefly warmed, and
examined under a compound microscope to find fungal structures inside
nematodes.

Results and discussion

Pinatubo Manandhar et Mew gen. nov. et P. oryzae Manandhar et Mew
sp. nov. (figs. 1-31).

Hypha sola hyalinum, laevibus, ramosis, rectis, septatum. Conidiophora
solitarius lata simplicia vel 1-2 ramosae, 1-4 septatis, hyalina, indeterminata
ad oryza seminibusque, cellae conidiogenae apice 0.7-2.0 um latae et
denticulae 0.7-1.3 x 0.6 um, plerumque 3-7 conidiis in capitulo laxum
ferentes. Conidiis solitarius hyalina, didymosporae; cellae aequalibus,
oboviformis usque piriformis usque clavatis, 6.7-10.2 x 2.2-3.3 um ad
substratis naturalibus, apice rotunda sensim adbasem tenuem, angusto
truncatoque fundamento minuentia. Sporodochiis conidioma, pionnotes
instratis, humido, pulvinata, persicino ochraceis, liberis ad aério hyphis in
oryza seminibusque. Conidiophora et cellae conidiogenae similis, 1-pluri
(plerumque 3-5)/conidiophora-mater-cella, lageniformis usque cymbiformis,
cum partapicem sporiferum flexuosa, capitulo laxum ferentes, muricatus vel
veruculosae et ventricoso rostrata, lecithiforme cum apice veruculosae.
Cicatrices conidiales conspicuae, saepe 0.3 x 0.3 um, primo terminates,
denique in positionem lateralem submotae et saepe plus minusve turgeo
prominentes. Conidiis sporodochis oboviformis usque piriformis usque
clavatis, sympodice producta, acrogenosa, minutulae denticulata,
amerosporae, raro 1-septatis, humidis et in massis persicino ochraceis, 5.6-8.4
x 1.8-2.6 um ad seminibusque, apice rotundatis sensim adbasem tenuem,
angusto truncatoque fundamento minuentia. Chlamydosporae intercalares in
hyphis, 1-2 mm densus, subter-aério mycelio et PDA superficies, sub-
globosae usque rectangularis.

204

Pinatubo Manandhar & Mew new genus and P. oryzae Manandhar & Mew
(figs. 1-31). This fungus is named after the legendary volcanic mountain Mt.
Pinatubo of the Philippines that was erupted last July 1991.

Fungi imperfecti, hyphomycetes, colony effuse with sparse solitary
conidiophores and pionnotes, mycelium partly superficial and partly
immersed, 65 mm diam in 7 days at 28°C, colorless with grayish powdery
solitary conidiophores and tan, papillate sporodochia on WA (figs. 24 b and
d); 31 mm diam in 7 days at 28°C, surface velutinous white, reverse peach
yellow due to dense chlamydospores on PDA (figs. 24 a and c); sparse to
floccose hyphae with sparse to dense solitary conidiophores and pionnotes
amphigenous on rice grains (figs. 25-27); mycelium superficial, hyaline,
smooth, branched, septate, 0.8-2.1 um across. Solitary conidiophores,
sparse to abundant in aerial mycelia on rice seeds and on PDA, straight from
lateral hyphae, macronematous, smooth, erect or suberect, simple or 1-2
branches, 1-4 septate (figs. 1, 26 and 29); sparse to numerous, branched
200-400 um tall on WA (Fig. 2); polyblastic, hyaline, indeterminate,
raduliforme, 21.2-84.8 um tall, base 1.6-2.7 um wide, near apex 0.7-2.0 um

205

Figure 1-23. Camera lucida illustrations of Pinatubo oryzae: 1. Simple to
lightly branched solitary conidiophores arising from an aerial hypha on the
natural substratum (infested rice seed) or on potato-dextrose agar (PDA);
2. A much branched solitary conidiophore arising from the superficial hypha
on the surface of 2% water agar (WA); 3. Subglobose sessile pionnote on or
above the natural substratum hung on its hypha; 4. Lateral view of a
superficial pionnote with heaping-up of sporodochial conidia on WA; 5. Close-
up view of a solitary conidiophore with an apical cluster of 2-celled solitary
conidia on the natural substratum or on PDA; 6-9. Denuded solitary
conidiophore apices, showing variations in length and arrangements of spore-
bearing spurs with more common capitate head on natural substratum (6),
dilated head often found on the natural substratum (7), and less commonly
found racemes apices on the natural substrata but common on the cultural
media (8 and 9); 10. Variations of solitary conidia formed on solitary
conidiophores; 11. A giant conidium producing secondary conidiophore and
conidia; 12. Catenate chlamydospores formed in older hyphae from PDA; 13.
Initial development of a sporodochium by thickening and branching of a
single hypha; 14. Development of conidiophore-mother-cells and
conidiophores; 15 and 16 Sporodochial conidiophore-mother-cells showing
variations in length, shapes and bearing the number of conidiophores; 17
and 18. Conidiophores showing flexuous conidiogenous cells in the terminal
portion formed on WA and on the natural substratum under wet
environment (17) and a warty or coralloid terminal portion formed under
drier environment (18); 19. Sporodochial conidia showing variations in shape
and size; 20-23. Successive development of the sporodochial conidia and
the terminal portion of the conidiogenous cell giving birth of first conidium
on a minute denticle on the top of a clavate-shaped conidiophore (20),
second conidium initiated on similar minute denticle by elongating the axis
just below the first conidial attachment displacing it to a side (21), third
conidium borne on new axis at the extremity of the second conidium
displacing the second conidium as before (22), and fourth conidium borne
similarly displacing the third conidium a side and by that time first and
second conidia dislodge depending upon their maturity and possibly the
force exerted by the tufts of developing conidiophores and sporodochial
conidial mass (23). Bar for Fig. 1 to 4 is scale A, for 12 is scale B, and for 5-
11 and 13 to 23 is scale C.

wide on PDA and on rice grains (figs. 5,.6 and 29), mostly 3-7 solitary conidia
in apical clusters with mostly capitate head (figs. 6 and 29) and dilated tip
(fig. 7) in natural substrate and few racemes types (figs. 8 and 9) on agar
medium. Most solitary conidia borne in apical clusters (figs. 5 and 29), some
borne in series of clusters or irregularly along spore-bearing apex on minute
subcylindrical 0.7-1.3 x 0.6 um denticles (figs. 7 to 9), oboviform to pyriform
to clavate, holoblastic, acropleurogenous, dry, 2-celled, equal celled, 6.7-10.2
x 2.2-3.3 um (average 8.2 x 2.5 um) on rice grains, 6.9-11.3 x 2.4-3.5 um
(average 9.8 x 2.9 um) on WA, apex rounded, apiculiform with minute

Figure 24 to 31. Photomicrographs of Pinatubo oryzae. 24. Colony
characteristics of the fungus on potato-dextrose agar (PDA) (a), 2% water
agar (WA) (b), reverse view on PDA (c) and on WA (d) grown in 9-cm diam
plastic Petri dish 7 days after incubation at 28°C and 12-hr photoperiod
(fluorescent light); 25. Growth of the fungus on a germinating rice grain in
10 days after incubation on blotter method; 26-27. Close-up view of the
fungus on the rice grain showing solitary conidiophores and conidia (arrows)
(26) and pionnotes (arrows) (27); 28. A washed pionnote showing

207

sporodochial conidiogenous cells and young conidia (arrows); 29. Solitary
conidiophores and conidia; 30. A heap of sporodochial conidia on a
sporodochium on WA; and 31. Chlamydospores. Bars in fig. 25 = 1 mm, in
figs. 26-27 = 8 mm and in figs. 28-31 = 50 um.

truncate base at the attachment end (fig. 10), larger than sporodochial
conidia. Solitary conidia germinating from the base of the proximal cells
and/or the near apex of distal cells. Giant conidia rare, 10-13 x 3-5 um, 1-2
septate, while attached to conidiogenous cells produce secondary
conidiogenous cells and smaller and smooth secondary conidia (fig. 11).
Sporodochial conidioma: Pionnotes on rice grains (figs. 3 and 27) and on WA
(figs. 4, 27 and 30) sparse, pulvinate, peach yellow, 47-128 um diam, stroma
absent or loose. Individual hyphae (fig. 13) thickened, branched, forming
ovate to clavate conidiophore-mother-cells (Figs 14 to 16), 4.5-7.5 x 2.3-4.5
um, branching 2-4 times in older sporodochia. Sporodochial conidiophores
not different from conidiogenous cells (figs. 15 and 16), tufts of
conidiogenous cells, short, abundant, 1 to many (mostly 3-5) per
conidiophore-mother-cell, lageniform to cymbiform, basal portion 6.6-14.4 x
2.3-3.6 um, with elongated, flexuous, non-septate, roughened with numerous
minute subcylindrical denticles mostly 0.3 x 0.3 um, terminal portion of the
conidiogenous cell 0-35 x 1-2 um in WA (figs. 17 and 28) but elongated 4.5-
8.2 um long or lecithiform with warts 2-4 um diam on rice grains (fig. 18).
Sporodochial conidia born holoblastically at the flexuous or warted tip of the
conidiogenous cell displacing older sporodochial conidia to one side,
dislodged when mature leaving a conspicuous flat to slightly raised conidial
scar (figs. 20 to 23), acrogenous, oboviform to pyriform, occasionally clavate;
wet, hyaline, rounded apex, minute truncate base, amerospores, 5.6-8.4 x
1.8-2.6 um (average 6.7 x 2.1 um) on rice grains and 6.2-8.5 x 2.1-3.0 um
(average 7.3 x 2.4 um) on WA, rarely 1-septate, produced sympodially on
minute denticles, peach yellow in mass (fig. 19), smaller than solitary conidia.
First sporodochial conidium formed on a minute denticle at the top portion
of the sporodochial conidiogenous cell as blown-out obovate cell, elongation
of the axis occurs with the production of the second sporodochial conidium
as a blown-out obovate cell just below the attachment of the first conidium
and so on. The first formed sporodochial conidium is detached before or
after the fourth sporodochial conidium is produced causing terminal portion
of the conidiogenous cell to be somewhat helicoid (figs. 20-23).
Chlamydospores, intercalary in hyphae, holothallic, thick-walled in a dense
mat, 1-2 mm thick in between aerial mycelium and surface of PDA, 6.3-11.8
um diam when subglobose, 8.2-15.9 x 3.5-6.0 um when rectangular to hour-
glass shaped, peach yellow in mass, yellowish individually (figs. 12 and 31).

Habitat: On and in seeds and germinating seedlings of rice (Oryza sativa L.)
grown in the Philippines

Type: Original living cultures were deposited in American Type Culture
Collection (ATCC 90547) and Centraalbureau voor Schimmelcultures (CBS
158.94)., and infected or infested dried rice seeds were deposited in the

208

University of Philippines at Los Banos (UPLB) Herbarium Deposition Number
10284, Laguna, Philippines.

Detection of Pinatubo oryzae in rice seeds: Two to 51% seeds
were infested with P. oryzae (table 1) among 20 seed lots (20 cultivars)
grown at the IRRI fields during the 1992 wet season, while 0 to 10% seeds

in the 1993 dry season. Seeds were infested with the fungus during both
dry and wet seasons, with a significantly (P = 0.05) higher percentage of
seed infestation during wet than dry season, however, the degree of
infestation was not significantly different (table 1). P. oryzae was found to
cause seed rot and weaken germinated seedlings showing yellowish gray
lesions on plumules.

Nematodes of A. besseyi were not parasitized by P. oryzae. No
constricting rings, loops, 3-dimensional networks or sticky knobs were found.

Table 1. Percent and degree of infestation due to Pinatubo oryzae on 40
seed lots grown at the IRRI farm during 1992 wet season and 1993 dry
season based on 400 seeds of 20 cultivars.

Detection of P. oryzae on seeds

Infestation Range Percent Degree@

1992 wet season 2-51 13 ad 4

1993 dry season 0-10 2b 3
LSDo 05 10 NS

4Based on the surface coverage with the fungal hyphae, where 0 = none to
9 = 100% cover.

ONumbers following different letters are significantly different by FLSD.

Identifying Clues for Pinatubo oryzae during routine seed
health test: The fungus abounds on rice at the IRRI central research farm

at Los Banos. The fungus was detected in 0 to 51% seeds in rice seed lots
produced by farmers in the Philippines. Record shows P. oryzae seed
infestation was first detected in 1982. However, no record of biological
control of A. besseyi using Arthrobotrys spp. in rice plants was found in

the Philippines. The fungus is also believed to conceal many other
pathogens from detection during routine seed health testing of the rice
seeds in the blotter tests.

P. oryzae can be identified from its sparse to dense white mycelial
cover on the seeds when viewed under 12x magnified objective (fig. 25), its
capitate arrangement of dry, powdery solitary conidia at the tips of the
conidiophores when viewed under 25x magnified objective (figs. 26 and 29);
and the presence of sparse creamy wet pionnotes hanging under a thin
cover of aerial hyphae (fig. 28), but which may turn sub-globose to irregular,
wet, light to pink on rice seeds on blotter in 7 days. Aerial conidiophores

209

may look like conidial heads of Sarocladium oryzae (Sawada) W. Gams & D.
Hawksw. and Verticillium luteo-album (Link:Fr.) Subram. under 12x magnified
objective. However, one can view characteristic triangular or cruciform
arrangement of solitary conidia in apical clusters of P. oryzae when surface
viewed under a 25x magnified objective to differentiate it from the rest of
the sporulating fungi. S. oryzae forms shiny, cream-colored heads and V.
luteo-album forms verticillate conidiophores in globose heads as seen under
a 25x magnified objectives. Pionnotes of P. oryzae are distinctly different
from pionnotes of Fusarium Link:Fr., being of coarse texture and
macroconidial origin (Nath et al. 1970). Pionnotes of P. oryzae may look like
pionnotes of Gerlachia oryzae (Hashioka & Yokogi) W. Gams, being similar in
texture and color. G. oryzae tends to form pionnotes in chains on the
surface of the seed and mostly without mycelial cover (Mia et al. 1985)
when viewed under the compound microscope. Conidia of G. oryzae are 2-
celled, sickle-shaped and larger than those of P. oryzae.

New genus Pinatubo is erected to place this fungus on the basis of
similarities in the nature of solitary conidiophores but dissimilarity in the
nature of sporodochial conidiomatal anamorphs in the same niche (on rice
seed or agar cultures) to the original or subsequently revised description of
Arthrobotrys. Sympodio-conidiation occurs on both solitary conidiophores
producing uniseptate conidia and sporodochial conidiophores producing
nonseptate conidia on P. oryzae. Separate names for the solitary
conidiophore anamorph producing uniseptate conidia and the sporodochial
anamorph producing nonseptate conidia is not necessary according to the
International Code for Botanical Nomenclature Articles 7.10 and 59.3.
Solitary conidiophore and sporodochial conidioma are synanamorphs of P.
oryzae according to Gams (1982). Two-celled conidia formed in the solitary
conidiophores of P. oryzae are smaller than the smallest conidia in the
genus Arthrobotrys, A. stilbacea with dimension of 10.5-14 x 4-4.5 um
(Meyer 1958), but has near identical dimensions on larger conidia forming
species of the genus Sporothrix Hektoen & Perkins. Conidial dimensions of
S. vizei (Berk. & Br.) de Hoog are 6-9 x 2.5-3.5 um, but they are 1-celled
formed in solitary conidiophores with 3-4 conidia per apical cluster (de Hoog,
1978). Similarities of the sporodochial anamorph and production of one-
celled sympodio-conidia are found in the genus Beauveria Vuill., but the
conidia in this genus are dry, globose to oval, and smaller (MacLeod, 1954).
Dimensions of 1-celled sporodochial conidia of P. oryzae are larger than the
largest conidia formed in the genus Beauveria, B. tenella (Delacr.) Siem.
which are mostly 2-4 x 1.5-2.5 um (MacLeod, 1954). Denticles on solitary
conidiophores of P. oryzae resemble fungicolous Eriomycopsis Speg. in
shape and size; and condial dimension of E. minuta Deighton & Pirozynski
which are mostly 7.5-14 x 1.3-1.8 um (Deighton and Pirozynski, 1972) is
close to the dimension of solitary conidia of P. oryzae but differs in the
shape. P. oryzae has two morphologically different anamorphs that do not
fit the combinations of features described for any known species of the
genera Arthrobotrys, Beauveria, Eriomycopsis or Sporothrix. We erected the
genus Pinatubo and named our fungus as P. oryzae _ because of
dissimilarities to the genus Arthrobotrys following the expanded generic

210

concept of Schenck et al. (1977) and Oorschot (1985) that has not been
described before or after 1977 and Eriomycopsis_ following the expanded
generic concept of Deighton and Pirozynski (1972) that has not been
described before or after 1972; by sympodially proliferating conidiogenous
cells on both solitary conidiophores producing macroconidia and sporodochial
conidiomata producing microconidia in gloeoid mass having more or less
conspicuous minute subcylindrical denticles. The fungus did not show any
affinities of forming hyphal constriction rings or loops, 3-dimensional
networks or sticky knobs to capture the seed-borne nematodes of rice, A.
besseyi that multiplied rapidly on the test fungus. P. oryzae was not
found to be fungicolous. We presumed that P. oryzae is adapted being
resident of rice panicles and seedlings, but its interaction with rice and other
microorganisms is a subject to study.

Three different conidia are found in the genus Fusarium, in which
septate pedicellate conidia (macroconidia) produced from sporodochia
bearing phialo-conidiogenous cells are larger than nonpedicellate conidia
(microconidia) produced from solitary phialo-conidiogenous cells, while
mesoconidia are produced. from solitary sympodio-conidiogenous cells
(Pascoe, 1990). Sporodochial conidia produced in P. oryzae are unicellular
and smaller than two-celled solitary conidia formed on the solitary
conidiophores and might be termed here as microconidia, while macroconidia
might be used for those produced on the solitary conidiophores. It is
believed that the ability of conidia to be detached by vibration and
dispersed by air current is apparently related to sympodio-conidiogenesis.
Solitary conidia of P. oryzae are dry, while sporodochial conidia are wet,
being disseminated by air or rain splash with equal reliability. This fungal
species demonstrating sympodio-conidia produced on the sporodochium
being wet spores (figs. 28 and 30). It is not known how mucilage is
produced in P. oryzae to prevent sporodochial conidia from being dry. It is
presumed that sporodochial conidia are forced to detach by underlying
numerous flexuous conidiogenous cells and developing sporodochial conidia
in situ (figs. 20 to 23), thereby a brief flow of cytoplasmic mucilage might
have occurred. Denticles are smaller on sporodochial conidiogenous cells
than on the solitary conidiogenous cells.

Conidial shapes of P. oryzae normaliy seen on the infested rice grains
are described above. Measurements were made in 1 M KCl. However,
conidial measurements in 1 M KCI were larger, from solitary conidiophores
mostly 7.6-10.6 x 2.6-4.2 um (mean 9.2-3.2 um) and from sporodochial
conidioma mostly 6.2-10.3 x 2.3-3.4.3 um (mean 7.8 x 3.2 um) with some of
them become obovate and somewhat constriction in the septum making
workers difficult to recognize fungal species (Mankau 1989).

Acknowledgment: Authors greatly indebted to Dr. Jack D. Rogers,
Department of Plant Pathology, Washington State University, Pullman, WA
99164, Dr. C. A. Shearer, Department of Biology, University of Illinois, Urbana,
IL 61801 and Dr. Walter Gams, Centraalbureau voor Schimmelcultures, 3742
SK Baarn, the Netherlands for reviewing the manuscript and inspection of

211

the fungus. Thanks are also due to E. G. Gonzales, P. G. Gonzales, C. C.
Huelma and S. D. Merca for technical details.

Reference cited:

Barron, G. L., and Davidson, J. G. N. 1972. Nematophagous hyphomycetes:
Arthrobotrys anomala sp. nov. Can J. Bot. 50:1773-1774.

Cooke, R. C., and Godfrey, B. F. S. 1964. A key to the nematode-destroying
fungi. Trans. Br. Mycol. Soc. 47:61-74.

Corda, A. C. F. 1839. Pracht-Flora europaeische Schimmelbildungen. Lipzig &
Dresden: G. Fleischer.

Deighton, F. C., and Pirozynski, K. A. 1972. Microfungi. V. More hyperparasitic
hyphomycetes. Mycological Paper 128, 1-110.

Dowsett, J. A., Reid, J., and Kalkat, R. S. 1984. A new species of Arthrobotrys
from soil. Mycologia 76:559-562.

Gams, W. 1982. Generic names for synanamorphs? Mycotaxon 15:459-464.

Haard, K. 1968. Taxonomic studies on the genus Arthrobotrys Corda.
Mycologia 60:1140-1159.

Hoog, G. S. de 1978. Notes on fungicolous Hyphomycetes and their relative.
Persoonia 10:33-81.

Jarowaja, N. 1970. The genus Arthrobotrys Corda. Acta Mycologica 6:337-
406.

MacLeod, D. M. 1954. Investigation on the genera Beauveria Vuill. and
Tritirachium Limber. Can. J. Bot. 32-818-890.

Mankau, R. 1989. Problems in taxonomic differentiation of nematode-trapping
fungi. J. Nematol. 21:572 (Abstr.).

McCulloch, J. S. 1977. New species of nematophagous fungi from Queensland.
Trans. Br. Mycol. Soc. 68:173-179.

Meyer, J. A. 1958. Appareil conidien de Trichothecium roseum Lk. ex Fr.,
Cylindrocarpon congonsis nov. sp. et Arthrobotrys stilbacea nov. sp. Bull.
Cociete Mycol. France 74:236-248.

Mia, M. A. T., Mathur, S. B., and Neergaard, P. 1985. Gerlachia oryzae in rice
seed. Trans. Br. Mycol. Soc. 84:337-338.

PANG

Nath, R., Neergaard, P., and Mathur, S. B. 1970. Identification of Fusarium
species on seeds as they occur in blotter test. Proc. Int. Seed Test. Assoc.
35:121-144.

Onofri, S., and Tosi, S. 1992. Arthrobotrys ferox sp. nov., a springtail-
capturing hyphomycetes from continental Antarctica. Mycotaxon 44:445-451.

Oorschot, C. A. N. van. 1985. Taxonomy of Dactylaria complex, V. A review of
Arthrobotrys and allied genera. Studies in Mycology, No. 26, 61-91.

Pascoe, |. G. 1990. Fusarium morphology |: Identification and characterization
of a third conidial type, the mesoconidium. Mycotaxon 27:121-160.

Schenck, S., Kendrick, W. B., and Pramer, D. 1977. A new nematode-trapping
hvand a reevaluation of Dactylaria and Arthrobotrys. Can J. Bot. 55:977-
985.

Southey, J, F. 1986. Laboratory Methods for Work with Plant and Soil
Nematodes. Ministry of Agr., Fish. Food, London, 202 p.

Zopf, W. 1888. Zur Kenntniss der Infections-Krankheiten neiderer Thiere und
Pflanzen. Nova Acta Leop.-Carol. Deut. Acad. Naturf. 52:314-376.

MYCOTAXON
Volume LX, pp. 213-224 October-December 1996

A PRELIMINARY CHECKLIST OF AGARICALES OF TURKEY

ERTUGRUL SESLI AND SAFFET BAYDAR

Faculty of Fatih Egitim (Education), Department of
Biology, University of Karadeniz (Black Sea)
Technical, 61335 Trabzon-TURKEY

ABSTRACT

This paper presents a complete list of the
members of the Agaricales identified by the
authors and other scientists since the beginning
of macrofungal studies in Turkey (1932). Species
listed here (table 1) belong to 17 families of
Agaricales. The collection of species newly listed
for Turkey by authors are deposited in the
Herbarium of the Faculty of Fatih EgGitim
(Education) of the Karadeniz (Black Sea) Technical
University.

INTRODUCTION

Turkey has avery divers flora, and many
studies have been carried out concerning the
phanerogamic component. However, the cryptogamic
flora has not been studied as extensively.

The main studies concerning the Agaricales in
Turkey were made by Pilat (1932, 1933), lLohwag
(1957, 1964), Selik (1965), Selik and Aksu (1967),
Oder (1982, 1986, 1988), Oner (1972), Karamanoglu
and Oder (1973), Kotlaba (1976), Siimer (1989),
Watling and Gregory (1977), Giicin and Oner (1982),
Glicin (1987, 1990), Isiloglu and Watling (1992),
Demirel and Isiloglu (1993), Gzttirk et al. (1990),
Asan and <Giicin (1990), Solak and Giicin (1990,
1992), Sesli (1993) and Baydar and Sesli (1994).

All of these studies focus on Turkish larger
fungi. There is no distinct study devoted to
Agaricales. In fact, there is no paper compiling
the Turkish Agaricales found by the _ scientists

214

listed above. This paper provides’ the First
summary of knowledge about the Turkish Agaricales.

Turkey has a mild and rainy climate. The
summers are warm and the winters, cool; nearly all
seasons are normally wet with mild temperature.
The climate during the spring, summer and autumm
is ideal for fungal growth.

MATERIALS AND METHODS

The specimens identified by the authors were
collected between 1990 and1994 in Trabzon Province
of Turkey. Standard collecting techniques were
used. At each collecting site some ecolocigal and
morphological properties were noted, and color
photographs of the specimens were taken using
Zenit 122 camera with a macro objective. Then the
specimens were taken’ to ’the® laboratory and
examined microscopically, and some chemical
reactions were performed. At the same time spore
prints were taken. The materials were dried and
jdentified according to Levetin et al. (1990),
Guevara et al. (1985), Hesler (1960), lLincoff
(1988), Marchand (1971-1986), Miller (1988), Moser
(1983), Phillips (1981) /'Ramsbottom {196574 1989)"
Rayner” (1979), “Singer “(Y982), Tulloss (1989,
1990. 1991), ‘Tulloss and |Vindgren (992) sa eine
materials examined by the authors are deposited in
the Herbarium of Fatih Egitim (Educatin) Faculty
of Karadeniz (Black Sea) Technical University in
Trabzon Province of Turkey.

RESULTS AND DISCUSSION

Species preceded by an asterisk (*) are new
records for Turkey which were found by the authors

and discussed previously (Baydar and Sesli, 1994;
Sesli, 1993)

In this study, 329 taxa are listed
distributed among families of Agaricales as
follows:

Tricholomataceae 34.34 % Agaricaceae 5.47 %
Boletaceae 10.94 $% Amanitaceae 5.47 %
Cortinariaceae 8.20 % Lepiotaceae 5.16 %
Coprinaceae 6.99 % Hygrophoraceae 3.95 %
Strophariaceae 5.47 % Pleurotaceae 3.34 %

Pluteaceae LIT GHAS
Bolbitiaceae 2.43 %
Crepitotaceae Diilerees
Paxillaceae LS se
Table 1. THE
BOLETACEAE

Boletus aereus Bull.:Fr.

B. aestivalis Fr.

B. calopus Fr.

B. chrysenteron Bull.

:Amans

B. edulis Bull.:Fr.

B. elegans Schum.

B. erythropus (Fr.:Fr.)
Pers.

B. luridus Schaeff.:Fr.

*B. porphyrosporus Fr.:
Hok.

B. queletii Schulzer

B. radicans Pers.:Fr.
B. regius Krombh.

B. retipes Bull.

B. rufus Schaeff.

B. satanoides Smotlacha
B. versicolor Rostk.
*B. zelleri Murr.

Leccinum carpini (R.
Schulz. )Moser

L. corsicum (Roll.)
Singer

L. insigne (Smith &
Thiers)Wat ling

*L. oxydabile (Sing. )

Sing.

L. quercinum (Pilat)
Green & Watling
L. scabrum (Bull.:Fr.)S.
F.Gray
L. subleucophaeum Dick &
Snell
L. versipelle (Fr.)Snell

Suillus bellini (Inz.)
Watling

S. boudieri (Quélet)
Watling

S. bovinus ( L.:Fr.) O.
Kuntze

Entolomataceae 0.91 %
Rhodophy 1 laceae 0.60 &
Gomphidiaceae 0.30 &

LIST OF THE SPECIES

S. granulatus (L.:Fr.)oO.

Kuntze
*S. pictus (Peck) Smith
& Thiers.

S. luteus (L.)S.F.Gray.

S. variegatus (Swartz:
Fr.)O.Kuntze

Trachypus pseudoscaber
Kallenb.
Tylopilus felleus (Bull.
TE) KRanecs

Xerocomus badius (Fr.:
Br.) Gal b:

X. subtomentosus (L.)
Quel.

GOMPHIDIACEAE
Chroogomphus rutilus
(Schaeff.:Fr)O.K.Miller

PAXILLACEAE
Omphalotus olearius
(DC.:Fr.)Singer
Paxillus atrotomentosus
(Batsch:Fr.)Fr.
P. involutus (Batsch.:
Fr.)Fr.
P. panuoides Fr.
P. rhodaxanthus Schw.

PLEUROTACEAR
Pleurotus cornucopiae
Paul .:Fr.
P. dryinus (Pers.:Fr.)
Kumm.
P. eryngii (D.C.:Fr.)
Quél.

P. ilgazicus Pilat

P. nemecii Pilat

P. ostreatus (Jack.:Fr.
)Kummer

216

P. ostreatus var.
salignus (P.:Fr.)Kummer
P. pubescens (Sow. )
Schroet.
P. pulmonarius (Fr. )Quel
P. serotinus var.
flaccidus Lange
P. ulmarius (Bull.:Fr.)
Quél.
COPRINACEAE
Coprinus atramentarius
(Bull.:Fr.)Fr.
C. comatus (Miiller :Fr.)
S.F.Gray
. disseminatus (Pers. )
Fr.
galericuliformis Losa
:Watling
micaceus (Bull.:Fr.)
Bx.
niveus (Pers.:Fr.)Fr.
ovatus Fr.
picaceus (Bull.:Fr.)
S.F.Gray
Cc. radiatus (Bolt.)Fr.
Lacrymaria lacrymabunda
(Bull.:Fr.)Pat.
L. lacrymabunda var.
velutina Pers.
Panaeolus acuminatus
(Schff.:Secr.)Quél.
P. campanulatus (Bull.:

Chek? tees ee pee)

Fr. )Quél.

P. fimicola (Fr.)Quel.
Ricken.

P. foenisecii (Pers.:

Fr.)Schroter
P. retirugis (Fr. )Quel
Psathyrella candolleana
(Fr.)Maire
P. fatua (Fr.)Konr.&
Maubl.
P. gracilis (Fr. )Quélet
P. melanthina (Fr. )Kihn.
& Romagn.
P. obtusata (Fr.)A.H.
Smith
P. prona (Fr.)Gillet

P. spadiceogrisea (Fr. )
Maire

PLUTEACEAE
Pluteus atricapillus
(Secr.)Sing.
P. cervinus (Schaeff.:
Fr. )Kummer
P. minutissimus R. Mre.
P. petasatus (Fr.)Gill.
P. salicinus (Pers.:Fr.)
Kummer
P. tomentosulus Pk.
Volvariella bombycina
var. maxima Pilat.
V. media (Schum.:Fr.)
Sing.
V. speciosa (Fr.:Fr)Sing

RHODOPHYLLACEAE
Rhodophyllus lanicus
Romagn.
R. sinuatus (Bull.:Fr.)
Sing.
ENTOLOMATACEAE

Clitopilus prunulus
(Scop.:Fr.)Kummer
Entoloma clypeatum (L.
:Fr.)Kummer
E. lividum (Bull.:St.
Amans )Quel.

HY GROPHORACEAE
Camarophyllius
subradiatus
(Schum. )Wiinsche
C. virgineus (Wulf.:Fr.)
Kars.
Hygrocybe conica (Scop.:
Fr.)Kummer
*H. langei Kiihn.
H. nigrescens (Quel.)
Kiihner
*H. subglobispora Orton
Hygrophorus calophyllus
Karst.
H. chrysodon (Batsch:
Fr.)Fr.

croceus (Bull.)Bres.
eburneus (Bull.:Fr.)
Fr.
fuscoalbus (Lasch)Fr.
pudorinus (Fr.)Fr.
russula (Schaeff.:Fr.
)Quél.

Lnrt tx

AMANITACEAE
Amanita vaginata var.
alba (De Seynes)Gill.

A. caesarea (Scop.: Fr.)
Quel.

A. citrina (Schaeff.)
Pers.

A. codinae (R.Maire)
Singer

*A. excelsa (Fr.:Fr.)
Bertillon

gemmata (Fr.)Gill.

junquillea Quélet.

muscaria (L.:Fr.)Pers.

A.
A.
A.
A. ovoidea (Bull.:Fr.)
Link
A. pantherina (DC.:Fr.)
Krombh.
A. phalloides Link:Fr.
A. spissa (Fr.)Kummer
A. rubescens Pers.
A. vaginata (Bull.:Fr.)
Viet
A. valens (E.J.Gilb.)
Bertault
A. verna (Bull.:Fr.)
Lamarck
A. virosa Lamarck
Torrendia pulchella Bres

AGARICACEAE

*Agaricus arvensis
Schaeff.:Secr.

augustus Fr.
bernardii (Quél.)Sacc
bisporus (Lange)Pilat
bitorquis Moll.
campestris L.:Fr.
comtulus Fr.
cupreobrunneus (Schff
-& Steer:Moell.)Moell.

*

a eo

217

lanipes (Moell. &
Sehft..) Sings
meleagris J. Schaeff.
rodmani Weck
semotus Fr.
Silvicola (Vitt.)Peck
xanthodermus Genevier
xerampelinus Schaeff.
A. variecolar Pers.
Psalliota edulis var.
valida Médllier
P. perrara (Schulzer)
Bres.

ee eS

LEPIOTACEAE
Cystoderma amianthinum
([Scop. ]Fr.)Fayod
Lepiota acutesquamasa
Gill
alba (Bres.)Sacc.
badhamii (Berk.Br. )
Quél.
brunneo-incarnata
Chod.:Martin
cristata Kummer
cygnea (Lange. )Mosser
holosericea (Fr.)Gill.
jubilaei Joss.
litoralis (Menier)
Boiffard

SSeS eas] S&S SS

L. naucina Fr.
L. rhodorrhiza Romagn.&
Locden P. DeOrtvon
Leucoagaricus
leucothites (Vitt.)
Wasser
Macrolepiota excoriata
Schaeff.:Fr.
M. mastoidea (Fr.)Singer
M. permixta Barla.
M. procera (Scop.:Fr.)

Sing.
BOLBITIACEAE
Agrocybe aegerita
(Brig.) Sing.

A. dura (Bolt.:Fr.)Sing.
A. semiorbicularis (Bull
.:St.Amans)Fayod

218

:Fr.)
Fay.
Bolbitius vitellinus
(Pers.:Fr.)Fr.
Canocybe lactea (Lange)
Metrod
C. coprophila (Kiihn. )
Kuhn.
C. rickeniana Sing.

A. pediades (Pers.

STROPHARTACEAE
Hypholoma fasciculare

(Huds.:Fr.)Kummer
Kuehneromyces mutabilis
(Schaeff.:Fr.)Sing &

Smith
Naematoloma

sublateritium (Fr.)Karst
Pholiota adiposa (Fr. )

Kummer
P. astragalina (Fr. )
Singer
P. aurivella (Batsch:Fr.
)Kummer.
P. caperate (Pers.:Fr.)
Kummer
P. destruens (Brond.)
Quel.
P. flammans (Fries. )
Kummer
P. radicosa (Bull.:Fr.)
Kummer
P. spectabilis (Fr. )
Kummer
P. squarrosa (Miiller:Fr.
)Kummer .

P. tuberculosa (Schaeff.
:Fr.)Kummer
Psilocybe pratensis P.D.
Orton

Stropharia aeruginosa
(Curtis:Fr.)Quélet
S. coronilla (Bull.:Fr.)

Quel.
S. semiglobata (Batsch.:
Fr.)Quel.

S. stercoraria Quél.

TRICHOLOMATACEAE
Armillaria bulbosa
(Barla:Kile)Watl.

A. mellea (Vahl.:Fr.)
Kummer.
A. tabescens (Scop.:Fr.)
Eme |
Clitocybe alexandri
(Gill)Konr.

C. cyathiformis (Bull.:
Fr. )Kummer

dealbata (Sow.:Fr.)
Kummer.

Q

ectypa Fr.
familia (Pk.)Sing.
fragrans (With.:Fr.)
Kummer
geotropa (Bull.:
Merat )Quél.
gibba (Pers.:Fr.)Quel.
gigantea (Sow.:Fr.)
Quel.
infundibuliformis
"(Schaef £. :Weinm. )Quél.
inversa (Scop.:Fr.)
Que].
laccata (Scop.:Fr.)
Kummer
lignatilis (Pers.:
Fr. )Kummer
perilis Gill
nebularis (Batsch:
Fr. )Kummer
odora (Bull.:Fr.)
Kummer
paropsis (Fries)Quel.
&Bres.
radicellata Gillet
rhizophora (Vel. )
Josserand: Pouchet
squamulosa (Pers.:
Fr.)Lange
tabascens (Scop. Meg ae
Gol lypie butyracea (Fr.
Bi) Rare
C. cessans Karst
C. dryophila (Bull.:Fr.)
Kummer

Q Cia aCe. eet, — eee nO) ae Cae) SSC) qa AO MA 0a

C. fusco-purpurea (Pers.
:Fr.)Kummer
C. grammocephala (Bull. )
FY
C. longipes (Bull.:St.
Amans)Mos.
C. peronata (Bolt.:Fr.)
sing:
C. tuberosa (Bull.:Fr.)
Kummer
Flammula hybrida (Fr.:
Fr)Gillet

F. liquiritiae Persoon.
F. penetrans (Fr.:Fr.)
Quel.
F. sapinea Quél.
F. spumosa (Fr.)Fr.
F. velutipes (Curt.:Fr.)
Karst.
Hohenbuehelia
atrocoerulea (Fr.:Fr. )
Sing.
(Kiihner )
Orton

H. rickenii

Laccaria amethystina Cke.

L. laccata (Scop.:Fr.)
Berk & Br.
Lepista luscina (Fr.:
Fr./S ing.
L. nuda (Bull.:Fr.)Cooke
L. saeva (Fr.)Orton
L. sordida (Fr.)Singer
Leucopaxillus amarus
(Alb. :Schw.:Fr. )Kiihn.
L. cerealis (Lasch)Sing.

L. gentianeus (Quél)Kotl.

Marasmius androsaceus (L

:Fr.)Fr.
M. calopus (Pers.:Fr.)
Exe.
M. caryophylleus
(Schaeffer)Schroet.
M. oreades (Bolt.:Fr)Fr.
M. perforans Hoffm.
M. ramealis (Bull.:Fr.)Fr.
M. resinosus Peck.
M. rotula (Scop.:Fr.)Fr.
M. splachnoides Fr.

219

Micromphale brassicolens
(Romagn.)Orton
Melanoleuca cinerascens
Reid
M. cognata (Fr.:Kon.)
Maubl. & Lge.
Mycena alcalina Kiihn.
M. atroalba (Bolt.:Fr.)
Gill.
corticola (Pers.:Fr.)
Quél.
epipterygia Bres.
fibula (Bull.:Fr.)
Kumm.
filopes (Bull.:Fr.)
Qué].
galericulata (Scop.:
Fr.)Gray
inclinata (Fr.)Quél.
integrella (Pers.:Fr.
)S.F.Gray
niveipes Murr.
parabolica Fr.
plicosa (Fries. )
Kummer
polygramma (Bull.)Fr.
forma pumila
pseudocorticola Kiihn
pura (Pers.)Fr. forma
rosea
sanguinolenta (Alb.&
Schw.:Fr.)Kummer
speirea (Fr.)Gill
stylobates (Pers.:
Fr.)Kumm.
strobilicola Favre &
Kiihner
subincarnata (Peck)
Sacc.
(Fr.)
Qué].
M. vulgaris (Pers.:Fr.)
Quel.
Omphalina abiegna (Bk.&
Br. }Singi.
O. gracillima (Weinm. )
Fries
O.hepatica(Fr.:Fr.)Gill.

x Kr KK K xx =

=

e

< £ £F KF EK

M. tintinnabulum

220

O. hydrogramma (Bull.:
Fr.)Quel.
O. leucophylla Fr.
O. velutina Quél.
O. olearius (D.C.:Fr.)
Sing.
Oudemansiella mucida
(Schrad.:Fr.)Hoehn.
O. radicata (Reh1l.:Fr)

Sing.

Panellus mitis (Pers.:
Fr.) Sing
P. stypticus (Bull.:Fr.)
Karst.

Xeromphalina campanella
(Batsch:Fr.)Mre.
Rhodocybe mundula
(Lasch) Sing.
R. truncata (Schaeff.:
Fra) Sindy
Strobilurus
stephanocystis (Hora)
L Sing.
Tricholoma albobrunneum
Heim
T. aurantium (Schaeffer:
Fr)Rick.
auratum (Paul.:Fr.)
Gillet
batschii Gulden
caligatum (Viv.)
Ricken
flavovirens (Pers.:
Fr.)Lund & Nanf.
grammopodium (Bull. )
Fre

ar |

- melaleucum Quél
orirubens Quél.
populinum Lge.
rutilans (Schaeffer. :
Ere) Sing.
sejunctum (Sow.:Fr. )
Quel.
(Fr.)Kummer
(Schaeff.:
Fr. )Kummer
T. virgatum (Fr.:Fr. )
Kumm.

HAS we A AA

%
ey

sordidum
terreum

a a |

CREPIDOTACEAE
Crepidotus cesatii Rab.
C. mollis (Schaeff.:Fr.)

Kummer

C. pubescens Bres.
*C. sphaerosporus (Pat. )
Lge.

Tubaria furfuracea
(Pers.:Fr.)Gill.

T. pellucida Bull.
T. trigonophylla (Lasch)
Fayod

CORTINARIACEAE
*Cortinarius
auroturbinatus (Secr.)

Lange.
*C. bulbosus (Fr.)Rick.
*C. bulliardii (Fr.)Fr.
C. cotoneus Fr.
C. malachioides P.D.

Orton
*C. glaucopus (Schff.:
Ershkhr,
Cc. olivaceo-fuscus
Kuehn.

Cc. punctatus (Pers.) Fr.
C. purpurascens Fr.

*C. subturbinatus Henry.
C. subvalidus R. Henry
Hebeloma leucosarx Orton
H. populinum Romagn.

H. sinapizans (Paulet:

Fr.)Gillet
Galerina marginata (Fr. )
Kuehn.

G. unicolor (Fr.)Sing.
Inocybe fastigiata
(Schaeff.:Fr.)Quél.
I. fibrosa var.
trivialis Lge.
I. geophylla var.
lilacina Gillet
gymnocarpa Kihn.
patouillardii Bres.
perbrevis (Weinm. )
Gill.
I. subrubens Atkinson.

et et

221

I. xanthodermus
Boursier & Kiihn.
Naucoria abstrusa (Fr. )

Fre
Phlegmacium amoenolens
RR: Hry.
P. variecolar (Fr.)
Wiinsche
ACKNOWLEDGMENT

The authors wish to thank Rodham E. Tulloss
for the identification of some species of Amanita
and reviewing the manuscript before publication.

LITERATURE CITED

Asan, A. and F. Giicin. 1990. Istiranca daglarinda
(Trakya) belirlenen bazi makrofunguslar. xX.

Ulusal Biyoloji Kongresi 1: 155-162.

Baydar, S. and E. Sesli. 1994. Trabzon ili
Akgaabat y6resinde belirlenen makromantarlar.
Doga-Tr. J. Bot. 18: 99-101.

Demirel, K. and M. Isilo@Glu. 1993. Ardanug (Artvin
) yOresi makrofunguslari. Yiiziinciti Y1l Universi-
tesi Fen Edebiyat Fakiiltesi Fen Bilimleri
Dergisi 4: 49-57.

Guevara, G., I. Garcia andJ. Castillo. 1985.
Algunos Agaricales del norte de Mexico. Rev.
Mex. Micol. 1: 129-188.

Gticin, F. and M. Oner. 1982. Manisa ili dahilinde
yetisen makrofunguslar. Doga Bilim Dergisi:
Temel Bilimler 6 (3): 91-96.

Giicin, F. 1987. Macrofungi of Piitiirge (Malatya)
in eastern Anatolia. Firat University 2 (1):
19-26.

Giicin, F. 1990. Elazig gevresinde belirJenen
makrofunguslar. Doga-Tr. J. Bot. 14: 171-177.
Hesler, R.L. 1960. Mushrooms of the great smokies.

Kingsport Press. Inc., U.S.A.

IsilogGlu, M. and R. Watling. 1992. Macromycetes
of Mediterranean Turkey. Edinb. J. Bot. 49 (1):
99-121.

Karamanoglu, K. and N. Oder. 1973. Bursa ili ve
cevresinde yetisen bazi gapkali mantarlar.
Ankara Universitesi Ecz. Fak. Mec. 3 (13): 13-
a35

222

Kotlaba, F. 1976. Contribution to the knowledge

of the Turkish macromycetes. Ceska Mycol.
30: 3-4.

Levetin, E., N. Jones, and K. Owens. 1990. A
preliminary checklist of the Agaricales of
Tulsa County, Oklahoma. Mycotaxon 36 (2): 337-
342.

Lincoff, H. G. L. 1988. The Audubon Society field
guide to North American mushrooms. Fourth
Edition. Chanticleer Press. Inc., New York.

Lohwag,), «Kk. 2957.3 PTUrK yernin mantar floras1
hakkinda arastirma. t.U. Orman Faktiltesi
Dergisi 7 (1): 128-137.

Lohwag, K. 1964. Belgrad ormanindan mikolojik
notlar. t&.U. Orman Fakiiltesi Dergisi 14 (2):
128-135.

Marchand, A. 1971-1986. Champignons du Nord et du
midi. tomes 1-9. Perpignan.

Miller, O.K. 1988. Mushrooms of north America.
Milton Rugoff Dai Nippon Printing Co., Tokyo.
Moser, M. 1983. Keys to agarics and boleti
(Polyporales, Boletales, Agaricales,

Russulales). London.

Oder, N. 1982. Kastamonu cevresinde yetisen
baz1i sgapkali mantarlar. S. WU. Fen Fakiltesi
Dergisi 2: 39-48.

Oder, N. 1986. Karadeniz bolgesinde (Sinop-
Artvin illeri arasi) yetisen onemli bazi
zehirli mantarlar uzerinde taksonomik

arastirmalar. S. U. Fen Edebiyat Fakiiltesi Fen
Dergisi 5: 87-104.

Oder, N. 1988. Karadeniz bolgesinde (Sinop-
Artvin illeri aras1) yetisen, halkin tanidigi
baz1 Onemli yenen mantarlar itizerine taksonomik
arastirmalar. S.U. Fen Edebiyat Fakiiltesi Fen
Dergisi 8: 215-236.

Oner, M. 1972. A contribution to the knowledge
of common Turkish higher fungi. Mycopathologia
et Mycologia applicata 47 (4): 369-373.

Ozturk; Ag (oDes Ha Anik ( andi/Ko Demirel. - °1990-

tnegol (Bursa) gevresinde yetigen zehirli ve
yenen mantarlar Uzerinde sistematik, morfolojik
ve ekolojik incelemeler. Yuzitincti Yil
Universitesi Fen Edebiyat Faktiltesi Fen

Bilimleri Dergisi 1 (1): 27-38.

Phillips, R. 1981. Mushrooms and other fungi of
Great Britain and Europe. First Edition. New
Interlitho S.P.A. Milan.

225

s

Pilat, Pew iO Site CONTE DUSONemEa ol etude des
hymenomycetes de l'’Asie Mineure. Bull. Soc.
Mycol. France 48: 162-189.

Pilat, A. 1933. Additamenta ad floram Asiae
Minoris hymenomycetum. Pars secunda:
Agaricineae. Bull. Soc. Mycol France 49: 283-
302e

Ramsbottom, J. 1989. Mushrooms and toadstools.
William SOtTLINe( "Sons ) feo) Coe Lt: Nineth
edition. London.

Ramsbottom, J. 1965. A handbook of the _ larger
British fungi. seventh Reprint. Alden Press.
Oxford.

Rayner, R. 1979. Mushrooms and toadstools. Hamlyn
Publishing Group Limited. Italy.

Selik, M. 1965. Belgrad ormaninda bulunan
yenilebilen mantarlar. t.U. Orman Fakiiltesi
Dergisi 15 (2): 48-55.

Selik, M. and S. Aksu. 1967. tstanbul'un park ve
korularindaki yerli ve yabanci agag _ tiirlerine
ariz olan odun tahrip eden mantarlar. tf.U.
Orman Fakiiltesi Dergisi 17 (1): 90-101.

Sesli, Eo 1993. Trabzon reg Pes Macka yoresi
makrofunguslari. Doga-Tr. J. Bot. 17:179-182.
Singer, (i Rai l962; The Agaricales in modern
taxonomy. Second Edition. Chr. Belser of

Stuttgart. Germany.

Sitimer, S. 1989. Some new records for the fungal
flora of Turkey. Marmara Universitesi Fen
Bilimleri Dergisi 6: 1-22.

Solak, M. H. and F. Giicin. 1990. Bursa yoresinden

baz} makrofunguslar. x. Ulusal Biyoloji
Kongresi 1: 163-171.

Solak, M. Bean dpaeGiue ling F. 1992. Bursa
Yoresinden Turkiye, t¢in Yeni Makrofungus
Turleri ve Yorede BelirJlenen Diger
Makrofunguslar. Doga-Tr. J. Bot. 16: 335-346.

Tulloss, R. E. 1989. Amanita eburnea - A new
species from Central America. Mycotaxon 36:1-7.

Tulloss, R. E. 1990. Amanita crenulata - history,
taxonomy, distribution, and poisonings.
Mycotaxon 39: 393-405.

buittose, Rk. EE. 1991. Amanita morrisii = history:

taxonomy, and distribution.Mycotaxon 40:281-286

Tulhoss, Rk. (Exand Js BE. ibindgrens 1992. .Amanita
smithiana-taxonomy, distribution, and poison-
ings. Mycotaxon 45: 373-387.

224

Watling, R. and M. N. Gregory. 1977. Larger fungi
from Turkey, Iran and neighbouring countries.
Karstenia 17: 59-72.

MYCOTAXON

Volume LX, pp. 225-240 October-December 1996

A NEW SPECIES OF THERMOASCUS WITH A
PAECILOMYCES ANAMORPH AND OTHER
THERMOPHILIC THERMOASCUS SPECIES FROM
TAIWAN

Kuei-Yu Chen* and Zuei-Ching Chen**

*Associate Professor, Department of Biology,
Chinese Culture University, Taipei, Taiwan, R.O.C.
**Professor, Department of Botany, National Taiwan
University, Taipei, Taiwan, R.O.C.

Abstract

A new thermophilic ascomycete isolated from field soils of Taiwan,
Thermoascus taitungiacus sp. nov. with a Paecilomyces anamorph is
described and illustrated. It is characterized by the presence of
predominantly echinulate ascospores under LM, but appearing
irregularly verrucose under SEM. The temperature range for mycelial
growth was between 25'C-50°'C. Abundant conidia and ascomata were
produced at the optimum growth temperature of 40°C and 30-35 °C,
respectively. No growth occurred at 20 C. Some other species of
Thermoascus from Taiwan are also discussed.

Key words: 7hermoascus taitungiacus, Paecilomyces, thermophilic
fungi, Thermoascus, soil fungi, Ascomycotina.

Introduction

During a survey of thermophilic fungi in Taiwan, three species
belonging to the genus 7hermoascus were isolated from field soils. One

226

of them different from the known species is proposed as a new species.
All dried culture specimens and living cultures are deposited in the
Mycological laboratory, Department of Botany, National Taiwan
University (TAI-Mycology). Capitalized color names are those in
Ridgway (1912).

Result and Discussion

Three species of thermophilic Thermoascus from Tatwan, 1.e.,
Thermoascus aurantiacus Miehe, T. crustaceus (Apinis & Chesters)
Stolk and a new species, 7. taitungiacus having Paecilomyces
anamorph are described as follows:

1. Thermoascus taitungiacus K. Y. Chen & Z. C. Chen sp. nov.

Fig. 1-3.

Coloniae in agaro" YpSs ", crescola totus catillus ad 40°C post six
dies. et aerius et decumbens hyphae, abundans agamicus structura cum
maxime efficiens aerius hyphae ac pauci granulae disseminatus. Collor
albesco ad Naples Yellow (Ridgway, 1912); reversum armeniacus flaius
cum aurantiacus flavus granulae.

Ascocarpae non ostiolatae, dispersae vel confluentes, subglobosae
vel irregulares, 330-600 um diam., flavae, aurantiacae vel brunneolae.
Asci glabri, hyalini, globosi vel subglobosi, octospori, (8.5) 10-13 x
10-15 um. Ascosporae galbanae, ovoideae vel ellipsoideae, globosae
vel subglobosae, 4.5-5.6 x 6.3-7.5 um, crassitunicatae, spinulosae.

Anam: Paecilomyces taitungiacus K. Y. Chen & Z.C. Chen sp. nov.

Conidiophora recta, hyalina, septata, glabra, phialides hyalina,
cylindrica vel ampullacea (2.8) 3.4-5.6 x 18-32 um, singulares vel
irregulariter 2-3 verticillatae. Conidia laevia, flavida, elliptica, (3-5 x 5-8
uum), vel subglobosa (2.5-6 tum diam).

Holotypus: Colonia exsiccata ex K-Y Chen 8709-2 (TAI-Mycology),
ver solum in " Antong " Taitung, Taiwan, 5.IX.1987.

Etymology: taitungiacus, referring to Taitung, the locality where this
new species was isolated in Taiwan.

228

Colonies on YpSs, growing all plate at 40°C in 6 days, both aerial
and prostrate hyphae abundant, asexual reproductive structures mostly
coming from aerial hyphae, mycelial granules scattered, White at first
becoming Naples Yellow, reverse Apricot Yellow with orange yellow
granules. At 35 C, colonies with abundant granules, covered by sparse,
erect aerial hyphae; the color Naples Yellow-Mustard Yellow,
Cadmium Yellow, then turning to Light Orange Yellow, reverse
Cadmium Yellow with Orange Rufous granules.

Ascomata nonostiolate, scattered or confluent, forming a crust-like
layer, covered by aerial hyphae and conidia, subglobose, or irregular in
shape, 330-600 um in diam., white at first, then yellow, light orange
yellow, orange, orange brown. Asci smooth, hyaline, globose or
subglobose mostly with 8 ascospores ( sometimes with 4-6 ascospores)
(8.5) 10-13 x 10-15 um evanescent at maturity. Ascospores yellowish
green, oval to elliptical, rarely globose to subglobose, 4.5-5.6 x 6.3-7.5
um, thick-walled, predominantly echinulate under LM, but irregularly
verrucose under SEM.

Conidiophores erect, arising from prostrate and trailing aerial hyphae,
hyaline, septate, smooth walled, consisting of an irregularly arranged
branch — system, and with the apex of each branch bearing 1-3
phialides. Phialides usually cylindrical or slightly flask-like, (2.8) 3.4-5.6
x 18-32 um, with a long conidium-bearing neck, up to 8 um long ,
single as side branches or appearing irregularly 2 or 3 at the end of
branches. Conidia in long chains at the tip of phialides, smooth, pale
yellow, cylindrical at first (2.2.-3.4 x 4.0-7.0 um), then elliptical
(3.0-5.5 x 5-8.0 um) or subglobose (2.5-6.0 tum in diam.) at maturity.

Specimen examined: TAIWAN: Taitung, Antong.. low land, weed soils,
5 IX, 1987. K-Y Chen 8709-2 (Holotype, TAI-Mycology).

Temperature tests: It is a thermophilic fungus. The optimum
temperature for mycelial growth was between 30°C and 40 C. The
most abundant production of conidia was observed at 40°C, followed at
30°C and 35 C, but ascocarp production was abundant between 30°C
and 35°C and very sparse at 40°C.

Note: The genus 7hermoascus was established by Miehe in 1907.

229

Fig. 2. Thermoascus taitungiacus sp. nov. (Chen 8709-2).
C. A portion of ascomal peridium. D. Asci. E. Ascospores in
evanescent asci. F. Ascospores.

230

Table 1. Radial growth, conidium and ascoma formation of
Thermoascus taitungiacus (Chen 8709-2) on YpSs
medium at various temperatures.

Temperature Mycelial growth Conidia Ascomata

Ke 6d 9d

*: Diameter of petri dish: 85mm

Growth: Colony diameter (mm) after 3, 6, 9 days (d).

Conidia and ascomata : (++++) very abundant; (+++) abundant;
(+) moderate; (+) slight and (-) not produced.

Several species of Thermoascus are thermophilic, including (7.
aurantiacus (Miehe, 1907); 7. aurantiacus var. levisporus
Upadhyay, Farmelo, Goetz & Melan (1984); 7. crustaceus (Stolk,
1965); 7. thermophilus (Sopp) von Arx (1970) and T. aegyptiacus
Ueda & Udagawa (1983). The taxonomy of these fungi is based
mainly on the morphology of their anamorph and teleomorph.
Thermoascus aurantiacus was first isolated by Miehe (1907),
followed by Apinis (1967). It was isolated from alluvial grassland soils.
Tansey (1973) also isolated it from alligator nesting material. They
had similar anamorph-aleurioconidia (a single, thick-walled, terminal
conidium); while no conidia were observed from those isolated by
Awao & Otsuka (1973); Minoura, et al.(1973); Stolk (1965) and Chen
& Chen (1988). Cooney & Emerson's isolate of 7. aurantiacus is most
likely identical with 7. crustaceus (Stolk, 1965). Upadhay, et al. (1984)
reported a new variety, 7. aurantiacus var. levisporus, which had an
anamorph similar to that of 7. aurantiacus. The anamorph of 7.
crustaceus ( Stolk, 1965) (=Dactylomyces crustaceus Apmis &
Chesters, 1967) and 7. aegyptiacus (Udea & Udagawa, 1983) were
confirmed to be the Paecilomyces, while T. thermophilus produced a
Polypaecilum anamorph. All species of Thermoascus had ellipsoidal

231

Fig. 3. Thermoascus taitungiacus sp. nov. (Chen 8709-2).
A. Ascomal initials. B & C. Ascospores. D. Conidia. E. Ascospores
(x 1250). F. Asci (x 1250). (A-D. Under SEM; E-F. Under LM).

232

and nearly smooth ascospores under LM. Under SEM, however the
ascospores appeared very minutely verrucose in 7: aurantiacus , finely
echinulate in 7. crustaceus, and verruculose in 7. thermophilus and T.
aegyptiacus. The ascospores of 7. taitungiacus were not only
predominantly echinulate under LM, but also irregularly verrucose

under SEM. It showed close affinity to 7. crustaceus m having a
Paecilomyces anamorph. Two kinds of conidia were observed in the
new species, one cylindrical or elliptical and the other globose to
subglobose. The latter usually developed at maturity.

2. Thermoascus aurantiacus Miehe, Die Selbsterhitzung des Heues,
p.70-73 (1907).

Fig. 4-5.

Colonies on PDA growing very rapidly, covering the plate at 40°C in
3 days, with erect white hyphae and granule-like ascomata, reverse
Dark Mars Brown.

Ascomata single or in groups forming a crust, first white, then turning
to yellow, orange, orange brown and dark liver brown, subglobose,
ovoid or irregular in shape, 280-560 x 370-672 um. Asci globose,
subglobose or ovoid, 8-11 x 10.6-14 wm with 4-8 ascospores
regularly arranged. Acospores elliptical, slightly roughened, 4-4.2 x
4.2-7 um. No asexual structure has been observed in our isolates.

Specimen examined: TAIWAN: Taipei, Tangsui Cheng, low land
cultivated soil of turnips. 20. IV .1984. K-Y Chen 8404-4. Pingtung
Hsien, Kueilai Hsiang, low land paddy soil, 10. VIII.1986. K-Y Chen
8608-28.

Temperature tests: Growth was very fast at 40°C and 50 C. No growth
occured at 30°C in 6 days. Ascomata were produced very abundantly
at 40°C and 50 C.

Note: Although Cooney & Emerson(1964) described the conidial
structure of 7. aurantiacus as Paecilomyces-like, no asexual structure
has been observed in our isolates. Other morphological characters of
our isolates are identical to the descriptions of this species reported by
Awo and Otsuka (1973).

30

Fig. 4. Thermoascus aurantiacus (Chen 8404-4).

A. Ascomal initial. B. Hyphae swollen near septum. C. Ascogenous
cells. D. Asci with ascospores. E. Ascospores in evanescent asci. F.
Ascospores.

234

Fig. 5. Thermoascus aurantiacus (Chen 8404-4).
A. Asci. B. Ascospores in evanescent asci. C. Ascospores. D.
Ascospores (x 1250). (A-C. Under SEM; D. Under LM).

235

Table 2. Radial growth and ascoma formation of
Thermoascus aurantiacus (Chen 8404-4)
on PDA medium at various temperatures.

Temperature Mycelial growth Ascomata
ag 3d 6d od 6d 9d

*:Diameter of petri dish: (92mm).

Growth: Colony diameter (mm) after 3, 6, 9 days (d).

Ascomata: (++++) very abundant; (+++) abundant;
(++) moderate; (+) slight and (-) not produced.

3. Thermoascus crustaceus (Apinis & Chesters) Stolk, Antonie van
Leeuwenhoek, 31: 272 (1965).
Anam: Paecilomyces crustaceus Apinis & Chesters, Trans. Br.
mycol. Soc. 47: 429 (1964).

Fig. 6-8.

Colonies on YpSs, growing very well, attaining all plate at 40°C in 3
days, bearing prostrate and aerial hyphae, granules scattered, covered
by erect hyphae, sometimes forming sectors. The color was white at
first, Antimony Yellow to Cadmium Orange; reverse Orange Chrome to
Orange Rufous. At 35°C, aerial mycelium sparse and granules more
abundant than those of 40 ic

Ascomata originating from ascogonial coils, nonostiolate, scattered
to confluent, mostly confluent forming a crusty layer, covered by aerial
hyphae.

Ascomata globose to irregular in shape 200-400 x 280-500 pm.
Color white at first, Yellow, Yellow orange, reddish orange. Ascomal
wall consisting of a few layers of pale yellowish brown
pseudoparenchyma cells. Asci single, hyaline, mostly 8-spored,
subglobose to pyriform, 11.5-12.5 x 13-16 um, thin-walled, evanescent
at maturity. Ascospores pale yellowish green, mostly elliptical, 4.5-5.1

500 oa

ae

Fig 6. Ther us (Chen 8504- fe
A.C ne sees ahd mhses s. B. Conidia. . D. Ascospores in
i. E. Ascospor

Z3T

x 6.0-7.5 um, rarely globose to subglobose, 4.5-5 um in diam.,
the surface with fine echinulations and some spores with 1-3 warty
structures.

Conidiophores septate, hyaline to pale yellow, smooth, arising
from prostrate and aerial hyphae, irregularly branched on the upper part
of conidiophores, measuring 3.5-5.5 x 6.8-36 tum, the apex of each
branch bearing phialides with a long conidium-bearing neck. Phialides
4.5-6 x 14.5-30 um, either solitary or in irregular verticils of 2 or
3 at the tip of branches. Conidia smooth, pale yellow, at first cylindric,
then elliptical, measuring 2.5-4.5 x 5-8.5 um, or sometimes becoming
globose or subglobose, 4.5-5 tim. in diam.

Specimen exammed: TAIWAN: Yunlin Hsien, Yunlin, low land
cultivated soils of peanut, 4.1V.1985. K-Y Chen 8504-10.

Temperature tests: The optimum growth temperature was between 30
°C and 40°C. Ascomata were very abundant between 35 C and 40°C,
abundant at 30 C. Asexual structures were very abundant at 30°C,
abundant at 35°C and 40°C.

Table 3. Radial growth, conidium and ascoma formation
of Thermoascus crustaceus (Chen 8504-10) on
YpSs medium at various temperatures.

Temperature Mycelial growth Conidia Ascomata
19 od 6d 6d

*: Diameter of petri dish : 85mm

Growth: Colony diameter (mm) after 3, 6, 9 days (d).

Conidia and ascomata: (++++) very abundant; (+++) abundant;
(++) moderate; (+) slight and (-) not produced.

238

Fig 7. Thermoascus crustaceus (Chen 8504-10).
Swollen branches bearing phialides, from culture at 40°C.

Note: Variations in phialide-bearing branches occurred at 40C,
including: (1) the width of the upper part of branches swelling up from
3.5-5.5 um at 30°C to 12 um ( 6-12 um) at 40°C (fig. 7) ; (2) the
number of phialides at the end of branches, increased up to 8 or more at
40°C, from 2 to 3 at 30 C; (3) mature conidia varied from elliptical at
30°C to globose at 40°C and (4) phialides becoming shorter at higher
temperature, i.e., from 14.5-30 pm at 30°C to 6-12 pmat 40°C. The
swelling of phialide-bearing branches at 40°C in our isolates was also
recorded for 7. aegyptiacus by Ueda & Udagawa (1983). The effect of
temperature on size of phialides was not mentioned by Stolk (1965).
However, this feature of the present isolate appear to be similiar to
Stolk's isolate (1965) at 30'C-35 °C and to Awao and Mistugi's isolate
(1973) at 40°C.

Acknowledgements

The authors are thankful to Drs. C. V. Subramanian and C. J. K.
Wang for reviewing this manuscript. This work was partly supported by

239

Fig 8. Thermoascus crustaceus (Chen 8504-10).
A. Ascospores in evanescent asci. B. Ascospores. C. Conidia. D. Asci
(x 1250). E. Ascospores (x 1250). (A-C. Under SEM; D-E. Under LM)

240

grants NSC 77-0211-B-002-01 and NSC 77-0606-B-002-167 to
Z.C.C. The technical assistances offered by the staffs of Electron
Microscope Laboratory, Department of Botany, NTU, are gratefully
acknowledged.

Literature cited

Apins, A. E. 1967. Dactylomyces and Thermoascus. Trans. Br. mycol.
Soc. 50: 573-582.

Arx, J. A. von. 1970. The genera of fungi sporulating im pure
culture. J. Cramer, Lehre, p 84.

Awao, T. and K. Mitsugi. 1973. Notes on thermophilic fungi in Japan
(1). Trans. mycol. Soc. Japan 14: 145-160.

Awao, T. and S. Otsuka. 1973. Notes on thermophilic fungi in Japan
(2). Trans. mycol. Soc. Japan 14: 221-236.

Chen, G. Y. and Z. C. Chen. 1988. Thermophilic and Thermotolerant
fungi in Taiwan (I). Trans. mycol. Soc. R.O.C. 3 (1): 1-72.

Cooney, D. G. and R. Emerson. 1964. Thermophilic fungi. W. H.
Freeman, San Francisco, 188 p.

Miehe, H. 1907. Die Selbsterhitzung des Heus. Eine biologische
Studie. Gustav Fischer, Jena. s. 127.

Minoura, K., K. Ochi and T. Nehira. 1973. Thermophilic filamentous
fungi in Japan (2). Trans. mycol. Soc. Japan 14: 362 -366.

Stolk, A. C. 1965. Thermophilic species of Yalaromyces Benjamin &
Thermoascus Miehe. Antonie van Leeuwenhoek 31: 262-276.

Tansey, M. R. 1973, Isolation of thermophilic fungi from alligator
nesting material. Mycologia 65: 594-601.

Ridgway, R. D., 1912. Color standards and color nomenclature. Publ.
by the author. Washington, D. C. 43 pp. 53 pl.

Ueda, S. and S. I. Udagawa. 1983. Thermoascus aegyptiacus, a new
thermophilic Ascomycete. Trans. mycol. Soc. Japan 24: 135-142.
Upadhyay, J. M., M. S. Framelo, S. O. Goetz and M. A. Melan. 1984.
A new variety of a thermophilic mold, Thermoascus aurantiacus

var. levisporus. Mycopathology 87: 71-80.

MYCOTAXON

Volume LX, pp. 241-247 October-December 1996

THIELAVIA PINGTUNGIA SP. NOV.
A THERMOPHILIC ASCOMYCETE FROM TAIWAN

Kuei-Yu Chen* and Zuei-Ching Chen**

*Associate Professor, Department of Biology,
Chinese Culture University, Taipei, Taiwan, R.O.C.
**Professor, Department of Botany, National Taiwan
University, Taipei, Taiwan, R.O.C.

Abstract

A new thermophilic fungus, Thielavia pingtungia sp. nov., isolated
from sugarcane field soilin Taiwan is described and illustrated. It is
characterized by brown, verrucose, hairy appendages on ascomata
and cylindrical asci with uniseriate, dark, ovoid to globose ascospores
with single apical germ pore. The optimum temperature for its growth

is between 35 C and 50 C.

Key words: Thielavia, Sordariaceae, Plectomycetes, Ascomycotina,
soil fungus, thermophilic, Fungal Flora of Taiwan.

During a survey of thermophilic fungi of Taiwan, an interesting
species of Thielavia with mouse-gray colonies and dark ascomata with
brown, hairy appendages was isolated. It has a high optimum growth
temperature range of 35°C to 50°C ; thus, it is considered to be
sufficiently different from other known thermophilic or thermotolerant
species of this genus ( Mouchacca, 1973; Pidoplichko, et al., 1973;
Lodha, 1974; von Arx, 1975a, b; Davidson, 1976; Moustafa, 1976;
Klopoteck, 1976; Udagawa & Sugiyama, 1981, 1982; Moustafa
& Abdul-Wahid, 1990) to warrant its description as a new taxon. All
dried culture specimens and living cultures are deposited at

242

Mycological Laboratory, Department of Botany, National Taiwan
University (TAI-Mycology). Capitalized color names are those in
Ridgway (1912).

Thielavia pingtungia K. Y. Chen & Z. C. Chen, sp. nov.

Fig. 1-2.

Coloniae in agaro YpSs ad 84 mm post 3 dies 40°C. Mycelia,
griseola. Hyphae 2.1-3.5 tum diam Color Murinus Canus ad Saturatus,
Murinus Canus, pallidus Fumus Canus in margo; reversum nigellus
Canus.

Ascocarpae nonostiolatae, superficiales, globosae vel
subglobosae 70-160 um, brunneae. Asci cylindrici, 39-52 x 7-9 um,
stipitati, octospori, hyalini vel flavidi, leptodermi. Ascosporae fere
uniseriatae, fere globosae vel subglobosae, brunneae, laeves,
crassitunicatae, 6.3-8.5 x 8.5-9.7 um.

Holotypus: Colonia exsiccata ex K-Y Chen 8608-5 (TAI-Mycology),
ager solum in " Changchin Hsiang", Pingtung, Taiwan, 10.VIII.1986.

Etymology: Pingtungia, referrmg to Pingtung, the locality where this
new species was isolated.

Colonies on YpSs, growing very fast, attaining 84 mm at 40°C in 3
days, aerial mycelium dense, composed of hyaline to pale gray,
septate, branched hyphae, 2.1-3.5 um in width, Mouse Gray to Deep
Mouse Gray and Pale Smoke Gray in the margin, reverse Blackish
Mouse Gray and Naple Yellow in the margin.

Ascomata bome on the hyphae, surperficial, non-ostiolate, globose
to subglobose, 70-160 um in diam., brown to deep brown or black
brown, composed of a pseudoparenchymatous wall, with flexuous,
undulate, brown, branched, thick-walled, hairy appendages, 2.5-4.0 um
in width, up to 350 um long. Asci cylindric, 39-52 x 7-9 um, stipitate,
fasciculate, 8-spored, hyaline to pale yellow, thin-walled, evanescent
at maturity. Ascospores usually uniseriate, few with an irregular
arrangement, globose to subglobose or ovoid, few elliptical and at one
end with a germ pore, at first hyaline, pale green, pale olive brown,
becoming dark brown with smooth and darkened thick-walls, some

243

Fig. 1. Thielavia pingtungia sp. nov. (Chen 8608-5).

A. Ascoma, B.A portion of ascomal peridium. C. Hair. D. Young
asci with undifferentiated ascospores, E. Asci with 8 ascospores. F.
Ascospores. G. Ascospores with one germ pore.

244

containing a large guttule, 6.3-8.5 x 8.5-9.7 (10.0) um. Conidia were
not observed.

Habitat: TAIWAN: Pingtung: Changchin Hsiang, Plain, Field soils of
sugarcane, 10 Vill, 1986. K-Y Chen 8608-5 (Holotype; TAI-Mycology).

Temperature tests: It is a thermophilic fungus. As shown in Table 1, the
optimum temperature is between 35°C and 50°C. Ascomata are in
general produced at 35 C and 40 C.

Thielavia pingtungia is a thermophilic fungus and differs from other
thermophilic or thermotolerant species of Thielavia (s. st.) by having
dark, septate, thick-walled, verrucose, branched, hairy appendages
uniformly distributed on the surface of ascomata. Most thermophilic
and thermotolerant species of Thielavia, i.e., T. aegyptica Moustafa &
Abdul-Wahid, 7. arenaria Mouchacca, T. australiensis Tansey & Jack,
T. emodensis Udagawa & Sugiyama, 7. expansa Udagawa &
Sugiyama, 7. heterothallica Klopotek, T. subthermophila Mouchacca,
T. terrestris (Apinis) Malloch & Cain,, etc. have smooth ascomata
and globose, ellipsoidal or clavate asci with irregularly arranged
ascospores. 7. microspora is a thermotolerant species with a weft of
brown, septate, branched, appendage-like hyphae, 1.5-3 um wide on
the surface of ascomata. Thielavia pingtungia is_ thus close
morphologically to T. microspora, but it differs in shape and size of asci
and ascospores. It is also quite distinct from other non-thermophilic
species of Thielavia having ascomata with hairy appendages or setae,
1.e., TJ. appendiculata Srivastava et al., T. fragilis (Natarajan) v. Arx,
T. hyalocarpa v. Arx, T. hyrcaniae Nicot, T. minuta (Cain) Malloch
& Cain, 7. peruviana (Gochenaur) Malloch & Cain, etc. T. pingtungia
is characterized by having cylindcric asci, 39-52 x 7-9 um, with
uniseriate, ovoid to nearly globose ascospores, 8.5-10 x 6.3-8.5 um. It
differs from species of Thielavia with cylindric asci, i.e., 7. tetraspora
(Lodha & Mirza) v. Arx with 4-spored asci, 7. tortuosa Udagawa &
Sugiyama with twisted asci, 7. octospora (Natarajan) v. Arx with nearly
sphaerical ascospores, 10-12 um in diam. and 7. hyalocarpa with
bigger asci, 70-100 x 11-18 pm.

245

Fig. 2. Thielavia pingtungia sp. nov. (Chen 8608-5).

A & B. Hairs on the surface of ascoma. (A. x 125; B. x 500), C. An
ascoma, D. Rough wall ofhairs, E-G. Ascospores, (F. x 500; G. x
1250), (A-B & F-G. Under LM; C-E. Under SEM).

246

Table |. Radial growth and ascoma formation of
Thielavia pingtungia (Chen 8608-5) on
YpSs medium at various temperatures.

Temperature Mycelial growth Ascomata
C 3d 6d Od

*: Diameter of petri dish: 85mm
Growth: Colony diameter (mm) after 3, 6, 9 days (d).
Ascomata: (+++) abundant; (++) moderate; (+) slight
and (-) not produced.

Acknowledgements

The authors wish to express their gratitude to Professor C.J.K.
Wang, Department of Environmental and Forest Biology, State
University of New York, College of Environmental Science and
Forestry, Syracuse, and Professor C. V. Subramanian, Department of
Botany, National Taiwan University, for reviewing the manuscript and
making helpful suggestions. This work was partly supported by grants
NSC 77-0211-B-002-01 and NSC 77-0606-B-002-167 to Z.C.C. The
technical assistances offered by the staffs of Electron Microscope
Laboratory, Department of Botany, NTU, are gratefully acknowledged.

Literature cited

Arx, J. A. von. 1975a. On Thielavia and some similar genera of
Ascomycetes. Stud. Mycol. C.B.S. 8: 1-29.

247

Arx, J. A. von. 1975b. On Thielavia angulata and some recently
described Thielavia species. Kavaka 3: 33-36.

Davidson, D. E. 1976. Thielavia pseudomaritima sp. nov. froma salt
lake in Wyoming. Trans. Br. mycol. Soc. 66: 175-178.

Klopotek, A.V. 1976. Thielavia heterothallica sp. nov., die perfekt
from von Chrysosporium. Arch. Microbiol. 107: 223-224.

Lodha, B. C. 1974. Studies on coprophilous fungi II. Thielavia. Nova
Hedwigia Beth. 47: 361-366.

Malloch, D. and R. F. Cain. 1973. The genus Thielavia. Mycologia 65:
1055-1077.

Mouchacca, J.1973. Les Thielavia des sols arides: Especés nouvelles et
analyse géenérique. Bull. Soc. Mycol. France 89: 295-311.

Moustafa, A. F. 1976. Thielavia kuwaitensis sp. nov. Trans. Br. mycol.
Soc. 66: 336-337.

Moustafa, A. F. and O. A. Abdul-Wahid. 1990. Thielavia aegyptica, a
new thermotolerant ascomycete from Egyptian soils. Persoonia 14:
173-175.

Pidioplichko, N. M., T. S. Kirilenko and V. A. Zakharchenko. 1973.
New species of the genus Thie(avia Zopt of the Ukrainian flora.
Mikrobiol. Zh. Kiev 35: 723-729 (in Russian).

Ridgway, R. D., 1912. Color standards and color nomenclature. Publ.
by the auther. Washington, D. C. 43 pp. 53 pl.

Udagawa, S.I. and Y. Sugiyama. 1981. Additions to the interesting
species of Ascomycetes from imported spices. Trans. mycol.
Soc. Japan 22: 197-212.

Udagawa, S. I. and Y. Sugiyama. 1982. New records and new species
of ascomycetous microfungi from Nepal, a preliminary report on
the expedition of 1980. On the Cryptogamic Study in Nepal
(Miscellaneous publication of the National Science Museum,
Tokyo), 11-46.

MYCOTAXON

Volume LX, pp. 249-256 October-December 1996

HYMENOSCYPHUS SEMINIS-ALNI,
A NEW SPECIES OF THE H. FRUCTIGENUS-COMPLEX *

Hans Otto Baral

Blaihofstr. 42, D-72074 Tiibingen, FRG

ABSTRACT

Hymenoscyphus seminis-alni is closely related to the H. fructigenus-
complex (Leotiaceae). It differs from the hitherto known members of this group
by long cilia on the ends of the ascospores, by the presence of croziers on the
ascogenous hyphae, and by spores partly septate already within the living asci.
The new species is so far only known from two localities in North and Eastern
Germany, occurring on dispersed previous year's seeds of A/nus in alder carrs.

ZUSAMMENFASSUNG

Hymenoscyphus seminis-alni steht dem H. _fructigenus-Komplex
(Leotiaceae) sehr nahe, unterscheidet sich von den bisher bekannten Sippen aber
durch lange Cilien an den Enden der Ascosporen, durch Haken an den ascogenen
Hyphen, sowie durch Sporen, die bereits im lebenden Ascus teilweise septiert
sind. Die neue Art ist bislang nur von je einem Fundort aus Nord- bzw.
Ostdeutschland bekannt. Sie besiedelt ausgefallene vorjahrige Samen von Alnus
in Erlenbriichen.

KEY WORDS: Ascomycetes, Leotiales, Leotiaceae, Hymenoscyphus
seminis-alni, Alnus seeds

In his contribution to the mycoflora of an alder carr near Bremen, Grauwinkel (1987:
62) presented a detailed description of a remarkable species of Hymenoscyphus Gray which
he collected on seeds of Alnus glutinosa, and which he tentatively named H. fructigenus
(Bull. : Fr.) Gray. Later, a further locality of this species was detected by M. Eckel near Taura
(Sachsen) on the same substrate. Through the kindness of both persons, I was able to obtain
fresh collections from both sites, allowing study of the micromorphology in vital state. These
specimens turned out to belong to an apparently undescribed species close to H. fructigenus.

* Contnbutions to the taxonomy of discomycetes V.

250

ABBREVIATIONS: H,0 = tap water, IKI = Lugol's iodine, KOH = potassium hydroxide 5%, CRB =
aqueous cresyl blue, * = living state (in HO), + = dead state, LB = lipid body, VB = refractive
vacuolar body, M = Botanische Staatssammlung Miinchen. Numbers in {} indicate in how many
collections the reported feature was tested and seen, or from which locality the feature derives.

Hymenoscyphus seminis-alni Baral, B. Grauwinkel & M. Eckel spec. nov.
Ethymol.: semen (lat.) = seed, Alnus = alder, referring to the substrate.

Apotheciis singulariter ex seminibus, disco 1-2.5 mm diam., + plano, albis vel pallide
ochraceis, glabris, stipitibus gracilibus, 6-20 x 0.15-0.45 mm. Ascis in statu vivo 120-140 x
8.7-10.5 tum, in statu emortuo 100-130 x 7-8 um, apice conico, IKI coerulescente, e uncis
natis. Ascosporis in statu vivo 16-24 x 3.5-5 um, in statu emortuo 15-23 x 3.2-4.5 yum, 0-1
septatis in asco vivo, rectis, raro leniter curvatis, apice rotundato, basi attenuato, cum paucis
guttulis magnis, polis cum 1-3 ciliis circiter 2.5-4.5 jum longis. Paraphysibus rectis,
cylindraceis, obtusis, in statu vivo multiguttulatis. Excipulo ectale textura prismatica (-
porrecta), strato externo hyphis guttulatis. Habitat ad seminibus emortuis Alni in Alnetis, in
sero autumno.

Holotypus: Germany, Bremen, Huchting, NSG Sodenmatt, 9.X1.1995, B.
Grauwinkel, HB 5370, depositus in M, 7 apotheciis.

Apothecia: disc 1-2.5 mm in diam. when fresh {2}, slightly concave to slightly
convex, white or very light cream, + brownish when dry, margin and exterior smooth,
concolorous, abruptly narrowed into a very long, cylindrical, often + undulating, concolorous,
sometimes somewhat transparent (glassy) stipe 6-20 mm long, 0.15-0.35 mm wide {2},
upwards widened to 0.4-0.45 mm and sometimes finely pubescent, base laterally inserted on
scale of seed, one apothecium per seed (very rarely two), no external sclerotinization, growth
negative geotropic. Asci *120-135 x 9.5-10.5 um {Taura}, *133-140 x 8.7-9.5 um
{Bremen}, tro (100) 115-125 x 7.2-7.7 um {Taura}, txow (100) 110-130 x 7-7.5 (8) um
{Bremen}, pars sporifera *50-67 uum long {2}, txow 68-85 tum, 8-spored, spores biseriate
(also in dead asci but here lower spores often + uniseriate), forcible spore discharge not rarely
seen in water mounts; apex distinctly (broadly) conical, apical thickening immature 1.7-2.4
jum, mature 1-1.3 um thick (tna), apical ring euamyloid, moderately to strongly blue (BB) in
IKI, occupying only the lower 1/3 - 3/4 of the thickening (Hymenoscyphus-type); base with a
+ short, somewhat undulating stalk, arising from croziers {4}. Ascospores ejected *16.3-20.3
x (4) 4.3-4.7 um {Taura}, *(14.3) 16-21 x 3.8-5 um {Bremen}, within living asci *16.5-20
(21.5) x 3.4-4 um {Taura}, within dead asci *(15) 16.5-20.5 (24) x (3.5) 3.8-4.7 um
{Taura}, tivo 16-20.5 x 3.5-4 um {Taura}, txow 15-23 x (3.2) 3.5-3.8 (4-45) um
{Bremen}, 0-1 septate within mature living asci (either all {Bremen} or c. 50% of the asci
{Taura} with 2-7 two-celled spores per ascus, especially the lower spores with septa),
straight, rarely very slightly curved, slightly to distinctly scutuloid, apex broadly rounded, with
a short lateral edge, moderately tapered towards + rounded base; both ends with 1-3 distinct,
mostly curved cilia (1) 2.5-4.5 (5) pm long {4}, cilia persistent in KOH (but easily
overlooked in that mountant); living spores with ~1-2 large LBs 1.3-2.3 am diam. (-2.8 um
through coalescence), surrounded by some smaller and several minute LBs (lipid content
comparatively high), LBs persistent in KOH, 1 nucleus ~3 um diam. in each cell; spore wall
surface + unstained in CRB. Paraphyses straight, cylindrical, *2.5-3.2 um wide near base

251

252

A) S| ke
@> r -@
pst 5 BC)
SP) e 'eNe
ok |e ae
ral ° A )
iS \O.i eg
_ A
33 ioral © (]
ist

ip

eo?
I ae ae ON,
6B0e

BS
eae

OO

a ,
as

—_

@x
XK

TOE BD,
Ee

SB
aes
Sal

Zc}

(DOOOR

OCYJo oO O°

DE CY
62S) CSREES verano

Rer = ae
5 Coleen:
Sere
~<a
@)

ie

©

11 12

253

(tkon 1.5-2.5 um), not or only very slightly enlarged towards apex, obtuse, protruding ~0-5
uum beyond asci (xox, mature living asci protrude 0-12 tm beyond paraphyses {2}), terminal
cell *40-74 x 3-4.2 (4.7) um {2} (ton 42-75 x 2.7-3.5 um), dichotomously branching
towards base and with many anastomoses; living paraphyses densely filled with strongly
refractive, globose, finally angular vacuolar bodies (multiguttulate, VBs) in the region from
apex (25) 40-80 (90) um downwards {2}, VBs deep turqueois in CRB, deep redbrown in
IKI, dissolving in KOH, disappearing in herbarium material (in dead paraphyses mounted in
H,0 the mentioned region may show a pale greyish-violet or yellowish-reddish plasmatic
colour through oxidation of a substance in the VBs). Ectal excipulum a horizontally oriented
(~10-20°), + undulating textura prismatica-porrecta, textura porrecta in stipe, on flanks ~40
um thick, individual cells *~15-45 x 11-18 um (tow ~20-50 x 6-13 pm), at base of
receptacle ~100 1m thick, cells *30-82 x (7) 10-17 (23) um {2}; covered by a single layer of
hyphae 3-6 um diam. containing scattered refractive guttules (VBs). Medullary excipulum:
inner part a + loose textura intricata, ~120 um thick near base of receptacle, of + undulating
hyphae *2-6 tm wide; outer part a dense textura porrecta, ~60-70 um thick near base of
receptacle, individual cells *42-82 x 3-4 (5) um, with anastomoses. Ecology: in late autumn
in alder carrs with /ris pseudacorus, Oenanthe aquatica, Alisma plantago, on previous year's
seeds of Alnus glutinosa {4} lying dispersed on or in wet litter below herbaceous vegetation.
A detailed description of the Bremen locality is given by Grauwinkel (1987: 4). The apothecia
are drought-intolerant except for the spores some of which survived for at least 2 weeks in
the herbarium.

Specimens examined (HB 4974b and HB 5370 in fresh state):

Germany: Bremen, Huchting, NSG Sodenmatt, MTB 2918, =2 m, 22.XI.1981, leg. B.
Grauwinkel, HB 2946; dto., 9.X1.1995, leg. B. Grauwinkel, HB 5370 (holotype, deposited in
M; isotype: herb. Grauwinkel). --- Sachsen, Taura, Kiihnhaider Wald, MTB 5043/3.3, ~320
m, 7.X1.1993, leg. M. Eckel, HB 4974a & herb. Eckel 93/31-07; dto., 14.X1.1993, leg. M.
Eckel, HB 4974b & herb. Eckel.

This new species so far only is known from two very remote sites of the northern and
eastern region of Germany. H. seminis-aini is distinguished from the common H. fructigenus
sensu lato (1) by the presence of long cilia on the spore ends, (2) by the asci arising from
croziers, and (3) by many or all of the living, mature, 8-spored asci containing 2-7 two-celled
spores. The long and tiny stalk might serve as a macroscopic character but is similarly present
in H. fructigenus var. carpini (Batsch) Hengstm. No other species of the genus is known to
me to occur on seeds of A/nus, an ecological niche to which the present species possibly is
adapted. The occurrence in late autumn also seems to distinguish it from H. fructigenus.

Figs. 1-15. Hymenoscyphus seminis-alni. Figs. 1-2. Apothecia on seeds of Alnus (fig. 1. x8, fig.
2. x3); figs. 3-5. Ascospores (x2000); fig. 6. Ascus with seven 1-septate spores (x400); figs. 7-8.
Ascus containing six aseptate and two 1-septate spores, paraphyses (x1000); fig. 9-10. Apices of
asci in IKI (x2500); figs. 11-13. Bases of asci with croziers (x2000); fig. 14. Section of
receptacle, x50; fig. 15. Section of ectal excipulum (lower flanks, x800). All from living cells in
HO except figs. 9-10 (IKI) and 13 (KOH). From Taura: HB 4974b: figs. 1, 4, 6, 12, 15; HB
4974a: 5, 9, 13; from Bremen: HB 2946: fig. 10; HB 5370 (holotype): figs. 2, 3, 7, 8, 11, 14.

254

Ciboria seminicola (Kienholz & Cash) Hechler (= Ciboria alni (Rostr.) Buchw.) also
occurs only on alder seeds but fruits in early spring (February). It is distinguished by hazel-
brown, cupulate apothecia of 2-6 mm diam. with stipes 2-12 mm long. 1-5 apothecia arise
from each seed. The apical apparatus of asci is typical for the Sclerotiniaceae to which this
species belongs, but the asci arise from simple septa, a rare feature in this family. The spores
are decidedly fusoid, *12-18 (21) x 4.5-5.5 uum, finely warted, without cilia, and contain only
some small LBs. The living paraphyses are devoid of distinct large guttules.

According to Grauwinkel (I.c.), the alder seeds infected by Hymenoscyphus seminis-
alni have the same size (2.5 x 2 x 0.5 mm) and colour (dark brown) as uninfected seeds, but
in section the embryo is absent, and the endosperm is comparatively thin and has an innermost
layer showing black stromatic colour. Those alder seeds infected by Ciboria seminicola are
distinctly larger (4 x 3.5 x 0.7 mm) and show a black stroma of the testa; the embryo is here
also absent while the endosperm is very abundant (see Grauwinkel, 1987: fig. 24).

Ciboria uliginosa (Fr.) Rehm as reported by Velenovsky (1934: 220) on cones of
Alnus in summer and autumn, is habitually similar to H. seminis-alni but is said to have
brownish apothecia and ellipsoid eguttulate ascospores 5-10 wm long suggesting a
Sclerotiniaceae. Authentic Ciboria uliginosa, however, is considered a synonym of
Cudoniella clavus (Alb. & Schw.: Fr.) Dennis (see Rehm, 1896: 1262).

The Hymenoscyphus fructigenus-complex is a very difficult group comprising several
fructicolous varieties or species. Each of these taxa is believed to be confined to fruits of a
different tree. All occur in early autumn ([Aug.] Sept.-Oct. [Nov.]) and not necessarily in wet
places like swamps etc. The type of H. fructigenus occurred on acorns of Quercus (Lizofi,
1992: 22). H. rokebyensis Dennis was separated on specific rank for collections on cupules of
Fagus. H. fructigenus var. aesculicarpa Sydow, H. f. var. coryli (Feull. in Rom.) Hengstm.
and H. f var. carpini were recognized for collections on cupules of Aesculus, seeds of
Corylus, and seeds of Carpinus respectively. | have examined a larger number of collections
on these substrates, also some on cones of Pinus, seeds of Acer, and seed scales of Juglans.
Collections on seeds of Prunus spinosa were reported by Dennis (1956: 80), on nut husks of
Hicoria by Seaver (1951: 121). "Helotium" nyssicola Seaver, on seeds of Nyssa sylvatica,
was restudied by Dennis (1964: 57); the ascospores are said to be without cilia, and Dennis
placed this taxon in synonymy of H. fructigenus.

A further apparently undescribed taxon of this complex was found by me to occur not
rarely on fallen previous year's cones of Alnus glutinosa (the cones did not contain any
seeds). This taxon differs by comparatively long [*(17) 20-28 (32) x (3) 3.5-4.5 (5) um],
distinctly scutuloid ascospores without or with 1-3 cilia (0.2-0.7 um long) at each spore end.
A further unidentified collection on this substrate differs from this and H. fructigenus by
short, thick, only slightly scutuloid spores (*14.5-19 x 4.5-5.5 um) without cilia.

The separation of this difficult complex of fruit-inhabiting species into different taxa
seems practicable but a thorough morphological investigation has not been undertaken. The
complex possibly is not a strictly monophyletic group because caulicolous, foliicolous, and
even lignicolous taxa seem very closely related. All collections of the H. fructigenus-complex
studied by me, and occurring on substrates other than A/nus seeds, differ consistently from H.
seminis-alni in at least two characters: (1) the asci always arise from simple septa, (2) the
spore ends are without cilia or have very short cilia (a collection on seeds of Corylus had 1-3
exceptionally long cilia [0.6-1.5 tum] at each spore end). Possibly, a third character: septate

253

spores within living asci, was also consistently absent in these collections (nevertheless,
septate spores may occur in these taxa outside the asci, or within dead asci). Finally, I have
not found any literature reports on such small seeds like those of A/nus as the host for species
of this complex.

Hymenoscyphus scutula (Pers. : Fr.) Phill. has cilia of a length similar as H. seminis-
alni, viz. (0.5) 2.5-3 (5) um, but there is only one cilium per spore end (rarely two). The asci
are consistently without croziers (see also White, 1942: fig. 6), and the apothecia have a
much shorter stipe. The species occurs on herbaceous stems of various dicots, also on large
petioles of deciduous trees. I tend to concur with White (I.c.) who found collections of H.
scutula with non-ciliate spores to be otherwise indistinguishable. Since I have studied only a
few specimens of H. seminis-alni, variation in both length and number of the cilia in this
species might similarly occur. In any case, cilia longer than c. 1 jm are a very rare feature in
the genus and have, according to my literature, so far only been reported for H. scutula (see
also Lizof, 1992: 10).

There exist, however, two seemingly undescribed species close to H. calyculus
(Sow.: Fr.) Phill. ss. auct., with 1-3 cilia at each spore end, 1-3 (4) um long: one occurs in the
Alps on twigs of Alnus (asci with croziers, spores *16-19.5 x 4.5-5.5 tum), the other was
found in Luxembourg on twigs of Carpinus (asci without croziers, spores *33-38 x 6.5-7
yum). In both species the large apothecia have comparatively short stipes (c. 0.3-1.5 mm
long), unlike H. seminis-alni.

The presence of long cilia in H. seminis-alni was considered by Grauwinkel (1987:
78) to facilitate spore dispersal in wet habitats. I cannot believe in this because the above-
mentioned species with ciliate ascospores usually do not occur in wet places.

The presence or absence of croziers is very often neglected in taxonomic studies on
the Leotiales. Thus, no information on this useful character can be found in the descriptions of
species of Hymenoscyphus in the studies of, e.g., Velenovsky (1934), Seaver (1951), Dennis
(1956), Svréek (1985), and Lizofi (1992). White (1942, 1943, 1944), however, carefully took
notice of the ascus base, depicted it accurately, and used it as an additional key character in
separating species of Hymenoscyphus sensu lato (as "Helotium"), e.g. in a key on page 139
(White, 1943). Dumont (1981 a, b), on the other hand, mentioned indeed the feature in his
descriptions of species referred to Helotium Pers. ex Gray and Hymenoscyphus, but he often
expressed his uncertainty about it, and he was unable to report a clear case of simple septate
ascogenous hyphae in any of the treated species although this case is so frequent in species of
Hymenoscyphus with scutuloid spores. Likewise, Huhtinen (1990) based the delimitation
within several closely related taxa of the genera Hyaloscypha Boud. and Hamato-
canthoscypha Svréek mainly on this character. In taxa he believed to be critical, Huhtinen
preferred to recognize only two different varieties, one with croziers, the other with simple
septa.

Dumont (1981 b: 80) found, "for some unexplained reason", a great variability in
spore septation when studying herbarium material of Hymenoscyphus caudatus (P. Karst.)
Dennis and allied taxa: in some collections all of the ascospores were 1-septate, while in
others most of them were aseptate and only few had a septum. One of the reasons for such a
variability is the fact that the number of septate spores strongly increases with the age of the
population. Contrary to this, the number of septate spores within living, mature
asci isamuch more constant value, being independent of the age of the apothecium, in my

256

experience, and having therefore a much higher taxonomical significance. This character
cannot, however, be re-examined from herbarium material since fresh apothecia usually
contain both living and dead asci, and all these asci are finally dead after desiccation (see
Baral, 1992: 376).

I prefer to recognize H. seminis-alni as different ftom H. fructigenus on specific level
because the deviating character of the ascus base is correlated with several other features
(cilia on the spores, spore septation, ecological and phenological differences).

ACKNOLEDGEMENTS

I wish to thank Bernt Grauwinkel (Bremen) and Matthias Eckel (Taura) for their help
in providing fresh specimens, and Prof. Donald H. Pfister (Cambridge) for reviewing a former
version of the manuscript.

REFERENCES CITED

BARAL, H. O. (1992). Vital versus herbarium taxonomy: morphological differences between
living and dead cells of ascomycetes, and their taxonomic implications. - Mycotaxon 44
(2): 333-390

DENNIS, R. W. G. (1956). A revision of the British Helotiaceae. - Mycol. Papers 62: 1-216

--- (1964). Remarks on the genus Hymenoscyphus S.F. Gray, with observation on sundry species
referred by Saccardo and others to the genus Helotium, Pezizella or Phialea. - Persoonia
3: 29-80

DUMONT, K. P. (1981 a). Leotiaceae II. A preliminary survey of the neotropical species
referred to Helotium and Hymenoscyphus. - Mycotaxon 12: 313-371

--- (1981 b). Leotiaceae III. Notes on slected temperate secies referred to Helotium and
Hymenoscyphus. - Mycotaxon 13: 59-84

GRAUWINKEL, B. (1987). Beitrag zur Pilzflora des Erlenbruchwaldes NSG Sodenmatt bei
Bremen. - Veréffentlichungen aus dem Ubersee-Museum Bremen A, 8: 1-165, pl. 1-2

HUHTINEN, S. (1990). A monograph of Hyaloscypha and allied genera. - Karstenia 29 (2):
45-252

LIZON, P. (1992). The genus Hymenoscyphus (Helotiales) in Slovakia, Czechoslovakia. -
Mycotaxon 45: 1-59

REHM, H. (1896). Rabenh. Krypt.-Flora, 2. Aufl., 1,3: Ascomyceten: Hysteriaceen und
Discomyceten. Leipzig

SEAVER, F. J. (1951). The North American cup fungi (inoperculates). New York, 428 pp.

SVRCEK, M. (1985). A taxonomic revision of Inoperculate Discomycetes described by J.
Velenovsky in the genus Helotium, preserved in National Museum, Prague ("1984"). - Sb.
Nar. Mus. Praze/ Acta Mus. Nat. Prag. (B) 40 (3-4): 129-215

VELENOVSKY, J. (1934). Monographia Discomycetum Bohemiae, Pragae.

WHITE, W. L. (1942). Studies in the genus Helotium. 1. A review of the species described by
Peck. - Mycologia 34: 154-179

--- (1943). Studies in the genus Helotium, Ill. History and diagnosis of certain European and
North American foliicolous species. - Farlowia | (1): 135-170

--- (1944). Studies in the genus Helotium, IV. Some miscellaneous species. - Farlowia | (4):
599-617

MYCOTAXON

AMOENODOCHIUM HUMICOLA ANAM. GEN. ET SP.
NOV., A NEW SPORODOCHIAL HYPHOMYCETE FROM
INDIAN SOIL

FERNANDO PELAEZ

Centro de Investigacion Basica
Merck, Sharp and Dohme de Espafia
Josefa Valcarcel, 38, 28027 Madrid, Spain

RAFAEL F. CASTANEDA RUIZ

Instituto de Investigaciones Fundamentales en Agricultura Tropical “Alejandro de
Humboldt”
Santiago de las Vegas, Calle 1, Esq. 1, Ciudad de la Habana, Cuba

Abstract

Amoenodochium humicola, isolated from soil in India, is described and illustrated as a
new genus and species of sporodochial hyphomycetes. The taxon is characterized by the
production of pulvinate green-brown sporodochia with sympodially proliferating
polyblastic and strongly denticulate conidiogenous cells, and turbinate or cuneiform one-
celled conidia, which are fimbriate at the base and crown-like at the apex.

Resumen

Se describe e ilustra Amoenodochium humicola como un nuevo género y especie de
hifomicetos esporodoquiales, aislado de suelo. El taxon se caracteriza por la produccién
en Cultivo de esporodoquios pulvinados verde-parduzcos, con células conidiogénicas
poliblasticas, proliferando simpodialmente, fuertemente denticuladas, y por conidios
unicelulares turbinados 0 cuneiformes, fimbriados en la base y ornamentados con una
corona en el pice.

Introduction

During a survey of tropical soil fungi, we isolated from a soil sample collected in
Goa, India, a fungus that developed green-brown sporodochia scattered on the agar
surface after a short period of incubation. When the isolate was examined
microscopically, the conidiogenous cells and conidia were seen to differ significantly
from all other known sporodochial hyphomycetes. A new genus and species is thus
proposed to accommodate this interesting fungus.

258
Materials and Methods

The fungus was isolated from a soil sample collected in a semi-deciduous forest
near Bondla, Goa, India, using a modification of the soil washing technique (Parkinson
& Williams, 1961), on DRBC medium (Oxoid). The growing colonies were transferred
to potato dextrose agar (PDA) plates for identification. Mature sporodochia were
observed after 7 days growth at 22°C under fluorescent light on a 12 hour light/dark
cycle. In the descriptions, capitalized color names are from Kornerup & Wanscher
(1963). PDA, oatmeal agar (OMA) and cornmeal agar (CMA) were prepared media
from Difco Laboratories. Yeast malt agar (YME) was prepared as medium M-67
described by Stevens (1981). Microphotographs were taken with a Leitz Diaplan
microscope equipped with Nomarski interference optics.

Description of new taxa
Amoenodochium Peldez et Castafieda Ruiz, anam. gen. nov.

Conidiomata sporodochia, superficialia, dissita vel confluens, pulvinata,
brunnea. Mycelium superficiale et partim in substrato immersum. Conidiophora
conspicua, mononematosa, fasciculata, geniculata, subhyalina usque pallide brunnea.
Cellulae conidiogenae polyblasticae, sympodiales, denticulatae, integratae, terminales.
Conidia turbinata, cuneiformia, late pyriformia usque obovoidea, unicellularia,
simplicia, sicca, brunnea, conspicua fimbriata ad basim. Teleomorphosis ignota.

Species typica: Amoenodochium humicola Peldez et Castafieda Ruiz, sp. nov.

Conidiomata sporodochial, superficial, scattered or confluent, pulvinate, brown.
Mycelium immersed and superficial. Conidiophores differentiated, mononematous,
fasciculate, geniculate, subhyaline to pale brown. Conidiogenous cells polyblastic,
sympodial, denticulate, integrated, terminal. Conidia turbinate, cuneiform, broadly
pyriform to obovoid, one-celled, simple, brown, conspicuously fimbriate at the base.
Teleomorph unknown.

Type species: Amoenodochium humicola Peléez & Castafieda Ruiz, sp. nov.

ETYMOLOGY: From the latin amoenus = "delightful", in reference to the
appearance of conidia and conidiogenous cells.

Amoenodochium humicola Peldez et Castafieda Ruiz, anam. sp. nov.

Mycelium ex hyphis septatis, laevibus, hyalinis. Conidiomata sporodochia,
superficialia, dissita vel confluens, pulvinata, brunnea, 100-500 tm diam.
Conidiophora conspicua, mononematosa, fasciculata, geniculata, levia, hyalina usque
leviter brunnea, 50-75 x 4-6 um. Cellulae conidiogenae polyblasticae, sympodiales,
denticulatae, integratae, terminales, hyalinae usque leviter brunneae. Conidia turbinata,
cuneiformia usque late pyriformia, unicellularia, simplicia, sicca, levia, subhyalina
usque pallide brunnea, conspicua fimbriata ad basim, coronata ad apicem, 7-14 x 4-7
iim, 2-3 um crassa ad basim. Teleomorphosis ignota.

TYPUS: Ex humus, Goa, India, 11 April 1994, F. Peldez. Teges culta sicca
(holotypus) in herbarium MA-Fungi (MA-33881). Isotypus in IMI.

Mycelium composed of septate, smooth, hyaline hyphae. Conidiomata
sporodochial, superficial, scattered or confluent, pulvinate, brown, 100-500 um diam.
Conidiophores differentiated, mononematous, fasciculate, geniculate, smooth, hyaline
to pale brown, 50-75 x 4-6 um. Conidiogenous cells polyblastic, sympodial,
denticulate, integrated, terminal, hyaline to pale brown. Conidia turbinate, cuneiform to
broadly pyriform, one-celled, simple, dry, smooth, subhyaline to pale brown,
conspicuously fimbriate at the base, crowned at the apex, 7-14 x 4-7 tum, 2-3 um wide
at the base. Teleomorph not observed.

259

Fig. 1. Amoenodochium humicola. A. Conidia. B. Sporodochium, conidiophores and
conidiogenous cells. Bar: 10 um.

260

ETYMOLOGY: From the latin humicola = "living in soil", in reference to the
habitat of the fungus.

HABITAT: from forest soil

SPECIMEN EXAMINED: Goa, India, 11 April 1994, F. Pelaéez. Dried culture
mat (holotype) in herbarium MA-Fungi (MA-33881). Isotype in IMI.

CULTURAL CHARACTERISTICS: On PDA attaining a diameter of 61-64 mm
in 2 weeks at 25°C and 50% RH, radially sulcate, margin uneven, appressed to -
submerged, aerial mycelium floccose, white. Sporodochia distributed in the inner two
thirds of the colony, eventually forming concentric rings, densely grouped at the center.
Colony Brownish Gray, dark Greenish Gray at the center, White at the margin.
Reverse Pale Yellow at the center, Yellowish Gray at the middle zone, Yellowish White
at the margin. Odors and exudates absent.

On CMA attaining a diameter of 48-52 mm, margin uneven, submerged, aerial
mycelium absent except for some sparse white radial strands. Colony colorless, with
few sporodochia scattered near the center. Odors and exudates absent.

On OMA attaining a diameter of 53-56 mm, margin uneven, appressed, aerial
mycelium sparse, subfelty, floccose, forming white radial strands. Brownish gray
sporodochia scattered over inner half of colony. Odors and exudates absent.

On YME attaining a diameter of 63-65 mm, radially sulcate, margin uneven,
appressed, aerial mycelium felty, floccose, Orange Gray (Alabaster), Grayish Orange.
Sporodochia absent, even after longer incubation. Reverse Light Yellow, Pale Yellow.
Odors and exudates absent.

The fungus grows optimally at about 25°C. At 22°C and 28°C the radial growth
rate decreases slightly with respect to the optimum (49-52 mm diam in PDA, in two
weeks). At 15°C growth was much slower (18-20 mm diam in the same medium and
period), and sporulation was absent. No growth was observed at 10°C nor at 37°C.

Discussion

Amoenodochium humicola shows some similarities to Asperisporium caricae
(Speg.) Maublanc, Polythrincium trifolii Kunze ex Ficinus & Schubert, (Ellis, 1971),
Scolecobasidiella tropicalis Castafieda & G. Arnold (Arnold & Castafieda, 1986) and
Ticogloea guttulata Weber, Spaaij & Gams (Weber et al. 1994), four hyphomycetes
associated with plants. All these species develop brown sporodochia or aggregations of
conidiophores resembling sporodochia, with sympodial conidiogenous cells. However,
the conidiogenous cells in genera Asperisporium, Polythrincium and Ticogloea are flat
and less conspicuous, whereas in Amoenodochium and Scolecobasidiella these are
conspicuously denticulate. The upper half of the branched conidiophores in
Scolecobasidiella terminate into sterile, filiform appendages, while in Amoenodochium
the conidiophores are entirely fertile.

The conidia of all five species are clearly distinct from each other. In A. caricae,
the conidia are almost always one-septate, ellipsoidal, pyriform or clavate, hyaline to
pale brown, verrucose and 14-26 x 7-10 um. In S. tropicalis, the conidia are one-
septate, brown, oblong, bluntly rounded at both ends, smooth, slightly constricted at
the septa and 8-12 x 2.5-4 um. The conidia in P. trifolii are one-septate, hyaline to pale
brown, oblong, cuneiform or pyriform, smooth and 17-24 x 13-24 um. In T. guttulata,
conidia are non-septate, dark brown, ovoid, slightly truncate at the base and

Fig. 2. Amoenodochium humicola. A. Two-week old culture mat on PDA (x 0.65). Note dark
sporodochia at the center. B. Sporodochia on OMA (x 4.2). C, D. Conidiophores and conidiogenous
cells with conidia attached (x 1250). E. Conidiophores and conidiogenous cells showing the denticles
(x 1250). F-I. Mature conidia (x1250). Note the fimbriate base (F, H) and the crown-like
ornamentation of the apex (G, I, top view).

262

4-5 x 3-3.5 uum. The conidia of A. humicola are one-celled, cuneiform, 7-14 x 4-7 um,
and patently fimbriate at the base.

The habitats of these fungi also differ. Asperisporium caricae inhabits living
leaves of Carica papaya L. in Central America, and P. trifolii is a widely distributed
plant pathogen causing sooty blotch disease on Trifolium. Scolecobasidiella tropicalis
was originally described from dead leaves of Calophyllum calaba L. in Cuba, but it has
also been found on leaf debris of cranberry (Vaccinium macrocarpon Ait.) in
Massachusetts, as a later synonym Mycosisymbrium cirrhosum Carris (Carris, 1994).
Ticogloea guttulata has been reported from roots of Ticodendron incognitum Gémez-
Laurito & G6émez P. and Tilia platyphyllos Scop., in Costa Rica and Germany
respectively. At present, Amoenodochium humicola is only known from soil and it has
not been isolated from leaf litter collected in the same area, although it cannot be
discarded that it may also be present in plant material.

Literature

- Arnold, G.R.W. & Castafieda Ruiz, R.F. (1986). Neue Hyphomyzeten-Arten aus
Kuba. Feddes Repertorium 97: 79-88.

- Carris, L.M. (1994). Vaccinium fungi: Mycosisymbrium cirrhosum gen. et sp. nov.
Mycologia 86: 131-133.

- Ellis, M.B. (1971). Dematiaceous Hyphomycetes. Commonwealth Mycological
Institute. Kew, England.

- Kornerup, A. & Wanscher, J.H. (1963). Methuen Handbook of Colour. Methuen,
London.

- Parkinson, D. & Williams, S.T. (1961). A method for isolating fungi from soil
microhabitats. Plant and Soil 13: 347-355.

- Stevens, R.B. (1981). Mycology Guidebook. University of Washington Press.
Seattle and London.

- Weber, G., Spaaij, F. and Gams, W. (1994). Ticogloea, a new genus of
hyphomycetes from roots of Ticodendron incognitum from Costa Rica. Mycol. Res.
98: 660-664.

MYCOTAXON

eee
__Volume LX, pp. 263-273 October-December 1996

MYCENA SORORIA SP. NOV., CLOSE TO
M. ROSEA GRAMBERG

(BASIDIOMYCOTINA)

J. PERREAU-BERTRAND, M.C. BOISSELIER-DUBAYLE
and J. LAMBOURDIERE

C.N.R.S. (U.MLS. 826, G.D.R. 1005)
Laboratoire de Cryptogamie, 12 rue Buffon. F - 75005 Paris.

Abstract - The electrophoretic comparison of six isozyme activities in Mycena
rosea and two differently pigmented - pink and lilaceous - forms of Mycena pura
(section Calodontes) demonstrates the specific characterization of MZ. sororia and
its affinity to M. rosea..

Key words - Electrophoresis, isozymes, biological species, taxonomy, Mycena,
Basidiomycotina.

Introduction

The electrophoretic analysis of isozymes provides some genetic
informations when morphological or other biochemical characters
appear to be insufficient for species delimitation. This technique is used
since a long time for fungal systematics (Blaich, 1977; Micales & al.,
1986; Molitoris & Prillinger, 1986) and many valuable results were
already obtained in several groups. Taxonomic units have been defined
by this procedure in diverse genera of Ascomycetes such as Morchella
(Yoon & al., 1990) and Tuber (Pacioni & Pomponi, 1991; Pacioni &

264

al., 1993) or Basidiomycetes : Polyporus (Shannon & al., 1973),
Pleurotus (Boisselier-Dubayle, 1983; Zervakis & al. 1994),
Armillaria (Lin & al., 1989) or Agaricus (Royse & May, 1982). For
the same purpose, a study was carried out, in the genus Mycena
(Tricholomatales), on M. pura (Pers. : Fr.) Kumm.

By the main features of its basidiomata, Mycena pura belongs
to the section Calodontes (Fr. ex Berk.) Quel. and more particularly to
the subsection Purae (Konr. & Maubl.) Maas Geest. In its orginal
sense, this taxon comprised four varieties differing from one another in
the colour of the pileus : pink, violaceous, purple and blue. Later on,
var. rosea was distinguished, on account of morphological criteria
(Maas Geesteranus, 1985, 1989), as a valuable specific entity named
M. rosea Gramberg. This distinction was corroborated by isozyme
patterns (Perreau & al., 1992).

At present time, 4 pura is considered to be a very variable
fungus (Comer, 1994) which is stated to be widespread throughout
northern and southern (Horak, 1979) temperate regions and some
tropical countries. In western Europe, seven colour forms were
described beside the epithet pura (Maas Geesteranus, 1989). Only
fruit-bodies which present a pale pink or lilaceous pink pileus and,
additionally, those having lilaceous, lilaceous grey or, sometimes, deep
violaceous cap, were examined for investigations on several isozymes
activities.

According to the electrophoretic data, the form with pink-
coloured pileus has proved to be genetically different from both the
lilaceous form on one hand and M. rosea on the other hand. These
results lead to the description of a new species, called Mycena sororia,
which appears more closely related to M. rosea than to M. pura sensu
lato.

Material and Methods

Material

One hundred twenty two basidiomata of Mycena rosea, 12 of
pink (representing MZ. sororia) and 29 of lilaceous or violaceous forms
of M. pura (29) were collected in Ile-de-France [Seine-et-Mame (77),
Essonne (91)] and the neighbouring departments of Aisne (02) and Oise
(60), during the autumns (October, November) 1992, 1993 and 1994.
The specimens were gathered in humus and leaf-litter under beech
(Fagus silvatica L.), beech mixed with oak (Quercus robur L.) or in
coniferous woods (Tab. 1) }

265

Locality Date Numbers of basidiomata
Forest or wood M. rosea WM. sororia M. pura

Fontainebleau (77) 1992 31 0 0
Fontainebleau (77) 1994 2 1 9
Fleury (91) 1993 0 0 5
Sénart (91) 1993 31 7 1
Hez (60) 1993 4 2 3
Coye (60) 1993 21 2 5
Coye (60) 1994 26 0 >
Laigue (02) 1994 i 0 1

Table 1 - Localities (names of the forests or woodlands) and number of
specimens collected for each locality and species of Mycena.

Methods

Field collected basidiomata used for electrophoretic studies had
reached maturity and were obviously at the same stage in their
development.

Isolates were prepared from explants of basidiomatal tissue.
Approximately 200-250 mg of tissue was ground with sand in 1 ml
tris-glycine buffer (4.95 mM, pH 8.3) at 4° C, with 0.34 %
mercaptoethanol, 1 % dimethyl sulfoxide, 7 mg polyvinyl pirrolidone
and 150 mg saccharose. The homogenate was centrifuged at 16 000 xg
for 15 min at 4° C and the supematant solution was removed and
stored at - 20° C before electrophoretic analysis (Boisselier-Dubayle &
Bischler, 1989).

The extracts were run in vertical acrylamide gels (2.5 %
stacking, 7 % migration) at 4° C, between glass plates 8 cm high and
0.75 mm thick (Hoefer apparatus), in a eae ert buffer (4.95 mM,
pH 8.3), with 10 mA per slab.

Of the ten enzymes surveyed, six aieioen active and well-
resolved bands. The enzyme staining solutions were prepared, just prior
to the end of each run, as described below.

Esterase (EST, E.C. 3.1.1.1) : 97 ml 0.05 M Tris-HCl buffer pH
7.1; 100 mg Fast Blue RR salt; 3 ml 1 % oP naphthyl acetate.

B glucosidase (GLU, E.C. 3.2.1.21) : 100 ml 6-bromo-2 naphthyl B-
D-glucopyranoside solution (10 mg 6-bromo-2 naphthyl B-D-
glucopyranoside; 20 ml methanol; 20 ml boiling water; 20 ml 0,2 M
phosphate-citrate buffer pH 4.95; 40 ml H,O); 20 ml 0.2 % tetra-
azotized O-dianisidine solution.

266

Acid phosphatase (ACP, E.C. 3.1.3.2) : 100 ml 0.2 M acetate buffer
pH 4.0; 50 mg a-naphthyl acid phosphate; 50 mg Fast Garnet GBC,
0.5 ml 10 % MgCl,.

Isocitrate dehydrogenase (IDH, E.C. 1.1.1.42) : 60 ml 0.1 M Tnis-
HCI - 0.01M MgCl, pH 7.5; 80 mg isocitric acid; 15 mg NADP; 6 mg
MTT; 3 mg PMS.

Glucose-6-phosphate dehydrogenase (G6PDH, E.C.1.1.1.49) : 100
ml 0.1 M Tris-HCl pH 7.0; 100 mg D-glucose 6 phosphate; 1 ml 1 M
MgCl,; 15 mg NADP; 15 mg NBT; 3 mg PMS.

Glutamate-oxaloacetate transaminase (GOT, E.C. 2.6.1.1) : 90 ml
0.1 M Tnis-HCl buffer pH 8.5; 100 mg a-ketoglutaric acid ; 200 mg
L-aspartic acid; ajusted to pH 8.5 with NaOH; 10 mg pyridoxal-5-
phosphate; 150 mg Fast Blue BB salt.

In the description of fresh material, reference is made to the
following, colour codes respectively indicated as :

K -_ Klincksieck, P. and T. Valette, 1908. Code des couleurs. Paris, P.
Klincksieck, 86 p.

M - Kornerup, A. and J.H. Wanscher, 1967. Methuen Handbook of
colour. London, Methuen & C°, 2nd ed., 244 p.

R -_ Rayner, R.W., 1970. A Mycological Colour Chart. Surrey, Kew,
Commonwealth Mycological Institute and British Mycological Society.

S - Séguy, E., 1936. Code universel des couleurs. Encyclopédie
Pratique du Naturaliste. Paris, Lechevalier, 48 pl.

Results

Electrophoretic patterns

The electrophoretic analyses of six isozyme activities show that
zymograms are differentiated into three main groups. They correspond
to three groups among the collected basidiomata : those with pink
pileus can be distinguished, on the one hand from those of the form
with violaceous or lilaceous pileus of Z. pura and, on the other hand,
from those related to M. rosea. Moreover, this latter species has been
previously separated from the complex M. pura, with the same methods
(Perreau & al., 1992). The samples with pink pileate basidiomata, until
now related to M. pura are named henceforth Mycena sororia.

Differences between the species of Mycena for three stainings
are illustrated in Figure 1. For EST and ACP, M. sororia exhibits a
very distinct zymogram. For IDH, the same two bands are revealed in

267

all individuals of MM. rosea (Boisselier-Dubayle & al., in press). Two
bands that always exhibit slower migration, are also observed in M.
sororia whereas only one band is revealed in M. pura.

Ss r r r Ss r F p p pp

Figure 1 - Electrophoretic patterns of Mycena sp. (s : M. sororia, r : M.
rosea, p : M. pura) for esterase (A), acid phosphatase (B) and isocitrate
dehydrogenase (C) activities.

A and B : samples of (4 sororia from Sénart are on the left, the others
came from Fontainebleau; C : the sample of 1Z. sororia came from Coye.

Electrophoretic changes are also observed for the three other
stainings. For GLU, one locus is revealed in the Mycena of the section
Calodontes. Two alleles are scored in M. rosea (Boisselier-Dubayle &
al., in press), whereas M. sororia appears monomorphic. In this
species, the allele exhibits a faster migration than those found in M.
rosea. As for GLU, only one locus is recorded for GOT; the
electromorph of M. sororia being of slower migration than those
detected in M. rosea and M. pura. G6PDH shows only one region of
activity : the electromorphs of M. sororia are nearly similar to those of
M. rosea but with a slower migration. Patterns of M. pura appear
different in mobility and intensity of bands.

268

Isozyme patterns of Mycena sororia appear therefore to be
specific, revealing nevertheless a certain affinity with MZ. rosea for IDH
and G6PDH. No noticeable variation was found among the
electrophoretic patterns of the 12 analyzed basidiomata. Indeed,
electromorphs of the specimens from the four forests and from two
years are identical.

Morphological characteristics

The specific status of Mycena sororia, well supported by
isozyme characterization, is corroborated by some macro- and micro-
scopical observations. The distinctive morphological characteristics of
M. sororia in comparison with M. pura are mainly the lilaceous to
whitish pink colour of the pileus and the larger size of the
basidiospores. Compared with M. rosea, the cylindrical or fusiform
outline of the stipe and the non-spheropedunculate cheilocystidia are
the most significant distinctive features. The shape and size of the
cheilocystidia, the presence or not of pleurocystidia are important
distinctive criteria also in other species of Mycenas of the subsection
Purae (Raithelhuber, 1980).

Description (Fig. 2)
Mycena sororia Perreau, Boisselier & Lambourdiere, sp. nov.

| Pileus 25-50 mm latus, campanulatus, convexus vel plano-
convexus, centro umbonatus, circum umbone concentrico-sulcatus,
translucente striatus, hygrophanus, lilaceo-roseus, pallidior, centro
flavescente. Caro tenuis, pallida, odore raphanoidei. Lamellae 38-42
stipitem attingentes, usque ad 5 - (7.5) mm latae, sinuato-adnatae,
albo-rosae, margine convexo, pallidiore. Stipes 70-95 x 4-6 mm,
cavus, cylindraceus vel fusiformis, levis, pallide roseo-albus deinde
lilacino-roseus vel purpureo-lilaceus, fibrillis sparsis basi instructus.

Basidia 30-35 x 5-6 um, clavata, 4-sporigera, fibulata,
sterigmatibus 6-7 longis munita. Sporae 7.5-8.5-10 x 4.5-5.5 jum,
inaequilateriter ellipsoideae, laeves, amyloideae. Cheilocystidia 50-
75 x 10-18 x 6-7 um, claviformia vel fusiformia, fibulata, levia.
Pleurocystidia similia, longiora, ad 20 jm diametro. Trama
lamellarum iodi ope vinescens. Hyphae pileipellis 2-5um latae,
fibulatae, leves. Hyphae stipitis corticales fibulatae, leves, cellulae
terminales 6-10 um latae, claviformae vel fusiformiae.

269

In nemore frondoso, ad terram, inter Fagus silvaticae vel
Quercus sp. folia dejecta.
Holotypus : J. Perreau , 94-339 (PC).

Figure 2 - Mycena sororia (Perreau, 94-339, holotype). Basidiomata - a :
habitus; b : longitudinal section; c : spores; d : cheilocystidia.

Basidiomata solitary or in small groups. Pileus up to 25-45-50
mm in diameter, campanulate to convex, then expanded, with
concentric depression around an obtuse conical - sometimes flattened -
umbo, shallowly rugose, translucent-striate, hygrophanous, lilaceous
pink [K 591/592; M 12 A 5; R 1, 58; S 15, 85, 664], "Aurora" pink,
whitish pink, pallescent with age [K 3 C, 53 C, 0121; M11 A 3; R61;
S 49, 254], often pale yellowish to yellowish pink at the centre; margin

270

concolorous. Flesh thin, whitish or light pinkish, with weak raphanoid
odour. Lamellae 38 - 42 reaching the stipe, tender, ascending then
almost horizontally, ventricose, up to 5 (7. 5) mm broad, sinuate-
adnate, dorsally intervenose, pale pink [K 578 B; S 70, 665], the edge
convex, sometimes slightly denticulate, whitish. Stipe 70-95 x 4-6 mm,
cylindrical or fusiform, not widened below, terete or a little
compressed, hollow, firm to brittle, smooth or finely striate, whitish
with pink and yellowish tint to lilaceous or purplish pink [K 28 A, 592;
M 11 A 3, 12 C 4; R 127; S 19, 55, 690], the base sometimes covered
with fibrils bringing together dead leaf-fragments.

Electrophoretically well characterized by EST, ACP, GLU,
GOT, IDH and G6PDH.

Basidia 30-35 x 5-6 um, clavate, tetra- (occasionally two- or
tri-) spored, clamped, with sterigmata up to 6-7 um long. Spores 7.5-
8.5-10 x 4.5-5.5 um, pip-shaped, smooth, amyloid. Cheilocystidia 50-
75 x 10-18 x 6-7 um, forming a sterile band (lamellar edge
homogeneous), smooth, clavate, subcylindrical , sometimes apically
broadly rounded or attenuated, fusiform, long-stalked, clamped.
Pleurocystidia numerous, similar, often longer (80-90 tum), up to 20
uum in diameter. Lamellar trama vinescent in Melzer's reagent. Hyphae
of the pileipellis 2-5 tum wide, clamped, smooth. Pigment vacuolar.
Hyphae of the cortical layer of the stipe 2-5 um wide, clamped,
smooth, terminal cells (caulocystidia) 40-100 x 6-10 um, simple,
claviform, fusiform, sometimes apically capitate, smooth.

In leaf-litter, under beech and/or oak (Fagus silvatica, Quercus
sp.), associated with M. rosea, but apparently less common. So far,
not found under coniferous trees.

Etymology : sororius, -a (soror), of sister, alluding to the affinity with
M. rosea.

Holotype : France, Seine-et-Mame, Foret de Fontainebleau, Route de la
Tour Deénecourt, 13. 11. 1994, J. Perreau N° 94-339 (PC).

Collections examined. France : Forét de Senart (91), 03. 10. 1993, J.
Perreau 93-200, 205, 206, 207, 208, 213, 214; Forét de Coye (60), 10.
10. 1993, J. Perreau 93-277, 279, Foret de Hez (60), 24. 10. 1993, J.
Perreau 93-389, 391.

Conclusion
271

Stable in several isozyme activities, M. sororia is rather
invariable in its morphological characters and in colour - pink with
lilaceous shade - of the pileus, only faintly yellowish or pallescent with
age and in dry conditions. It may partly represent f. pura of M. pura
described with "concentric depression round the umbo" of the pileus
(Maas Geesteranus, 1989). It is also not unlike to f. roseoviolacea
(Gillet) Maas Geest. of the same taxon, a form that has a pileus with
dingy or violaceous tints. Nevertheless, the species studied here can not
be assigned to M. pura, often described with lilaceous, grey lilaceous,
violaceous or purple basidiomata and smaller spores (6.2-7 x 3-4 um),
growing not only in deciduous forests but also under coniferous trees.

Mycena sororia has biochemichal affinities to MZ rosea, an
enzymatically well defined species with distinctive morphological
characters as stated by Maas Geesteranus. Often found in the same
habitat, probably mistaken for MZ rosea when it is not named M. pura,
M. sororia differs from the first taxon by the more violaceous pink
colour of the pileus which is also more dry in touch, and by the pinkish
white to purplish pink stipe, cylindraceous or fusiform, not widened at
the base; moreover it presents spores which are somewhat larger (7.5-
8.5-10 x 4.8-5.5 um) than those of M. rosea (6.5-9 x 4.5-5 Bioanal
spheropedunculate cheilocystidia.

On the other hand, the relationships between M. sororia and
the caespitose M. auroricolor (Berk. & Br.) Petch described from Sn
Lanka (Pegler, 1986) and mentioned as a member of the subsection
Purae (Maas Geesteranus, 1992) are unknown. The habit and colour of
the basidiomata seem similar in both M. sororia and M. auroricolor.
Nevertheless, in this latter species, the spores appear to be smaller and
pleurocystidia are absent. A comparison of fresh collections would be
required for a better separation of these two Mycenas originating from
geographically distant countries.

Two biological taxa inside Mycena pura with pink pileus are
therefore distinguished. MZ pura sensu lato, with its wide range in
colour and diversity in microscopical characteristics, has still to be
regarded as a complex of morphologically similar species. A revision of
this artificial entity and of its world distribution is now desirable. In
particular, further investigations are needed to test identity of

272

specimens from temperate and tropical origin (Comer, 1986, 1994;
Maas Geesteranus & Horak, 1995).

Acknowledgements
The authors express their gratitude to Dr. H. Bischler (C.N.R.S.,
Museum, Paris) for helpful suggestions and improvements in writing this
paper and to Prof. Dr. E. Horak (E.H.T., Zurich) for constructive comments
and critically reviewing the manuscript. They also thank M. Dumont-
Vialatte (Museum, Paris) for the photographic prints.

LITERATURE CITED

Blaich, R., 1977. Enzymes as an aid in taxonomy of higher
basidiomycetes. In : The Species Concept in Hymenomycetes
(Clémengon, H., ed.). Proc. Herbette Symp., Lausanne,
Switzerland, 1976. - Biblioth. Mycol. 61 : 215-228.

Boisselier-Dubayle, M.C., 1983. Taxonomic significance of enzyme
polymorphism among isolates of Pleurotus (Basidiomycetes) from
umbellifers. Trans. Brit. Mycol. Soc. 81 121-127.

Boisselier-Dubayle, M.C. and H. Bischler, 1989. Electrophoretic
studies in Marchantia polymorpha L. J. Hattori Bot. Lab. 67 : 297-
ay

Boisselier-Dubayle, M.C., J. Perreau-Bertrand and J. Lambour-
diere, in press. Genetic variability in wild populations of Mycena
rosea Gramberg (Basidiomycotina). Mycol. Res.

Corner, E.J.H., 1986. The tropical complex of Mycena pura. Bot.
Soc. Edinburgh, Transactions, 150 th Anniv., suppl. : 61-67.

Corner, E.J.H., 1994. Agarics in Malesia. I Tricholomatoid - I
Mycenoid. Nova Hedwigia 109 : 196-200.

Horak, E., 1979. Fungj, Basidiomycetes, Agaricales y Gastero-
mycetes secotioides. Fil. Criptog. Tierria del Fuego 11 : 1-525.

Lin, D., M.T. Dumas and M. Hubbes, 1989. Isozyme and general
protein patterns of Armillaria spp. collected from the boreal
mixedwood forest of Ontario. Canad. J. Bot. 67 : 1143-1147.

Maas Geesteranus, R.A., 1985. Studies in Mycenas 148-167. Proc.
Kon. Ned. Akad. Wetensch., C 88 : 47-62.

Maas Geesteranus, R.A., 1989. Conspectus of the Mycenas of the
Northern Hemisphere s 13. Sections Calamophilae and Calodontes.
Proc. Kon. Ned. Akad. Wetensch., C 92 : 477-504.

Maas Geesteranus, R.A., 1992. Some Myceneae of the Himalayan
foothills. Persoonia 15 : 33-53.

273

Maas Geesteranus, R.A. and E. Horak, 1995. Mycena and related
genera from Papua New Guinea and New Caledonia. Biblioth.
Mycol. 159 : 143-229.

Micales, J.A.. M.R. Bonde and G.L. Peterson, 1986. The use of
isozyme analysis in fungal taxonomy and genetics. Mycotaxon 27 :
405-449.

Molitoris, H.P. and H. Prillinger, 1986. Isoenzyme spectra for
characterization and identification of fungi (influence of genetical
and nongenetical factors). Mycol. Helvet. 2 : 103-122.

Pacioni, G. and G. Pomponi, 1991. Genotypic patterns of some Italian
populations of the Tuber aestivum - T. mesentericum complex.
Mycotaxon 42 : 171-179.

Pacioni, G., G. Frizzi, M. Miranda and C. Visca, 1993. Genetics of a
Tuber aestivum population (Ascomycotina, Tuberales). Mycotaxon
47 : 93-100.

Pegler, D.N., 1986. Agaric Flora of Sn Lanka. Kew Bull., Addit. Ser.
12: 1-519.

Perreau, J., J. Lambourdiére and M.C. Boisselier, 1992. Mycena
rosea et le complexe Mycena pura. Cryptogamie, Mycol. 13 : 247-
Zep

Raithelhuber, J., 1980. Der Arten- und Formenkreis um Mycena pura
(Pers. ex Fr.) Kumm. Metrodiana 9 : 26-40.

Royse, D.J. and B. May, 1982. Use of isozyme variation to identify
genotypic classes of Agaricus brunnescens. Mycologia 74 : 93-102.

Shannon, M.C., S.K. Ballal and J.W. Harris, 1973. Starch gel
electrophoresis of enzymes from nine species of Polyporus. Amer.
J. Bot. 60 : 96-100.

Yoon, C.S., R.V. Gessner and M.A. Romano, 1990. Population
genetics and systematics of the Morchella esculenta complex.
Mycologia 82 : 227-235.

Zervakis, G., J. Sourdis and C. Balis, 1994. Genetic variability and
systematics of eleven Pleurotus species based on isozyme analysis.
Mycol. Res. 98 : 329-341.

wr rr. re
A
‘ i
{ ae |
rT 1
2 2, a ey H
4s my ot Ke ‘ uy
1 } * ey me
P ‘ ps
vis ‘ ’ roe
p ;
4h 4 oF ray 14? aH Ply
- *
‘ Vi ey
by ’ j .
2 P 4 4
74 ae ¥
\
. i & 7
é : Ves ae |!
; :
|
“ nity n 4 f
2 1 ' 1
a - 2]
' \ Ra yt : E ce tine tk j
‘ , 4,
y ie LA a he av i Aid
, j + . ‘, i
yok { Ti “44 }
i ' ‘ 13 ed é
' i i 1 i y
a
t , hs .
: ‘ whl ; (
j “ld
vt rl ch . Peay 2 f
‘ f 4 as ,
7 \ . Te ;
| Nad at ee
} a. Ni Pe °
4 A ‘3 4 © * ib 7 (| 3
f >
\ [7
F te)
é
i
; ; tA, 4 ,
:
j 1%
}
Ge f j ; ‘
/ ‘
if
{ } \ / ‘ ; |
J u 4
a iY ; ’
es 4
4 ‘ ; r
" y j i : a
neg { , 4
%* 1 ara ¢ %
¢ } A ‘i
/ nn
* m 1 \ 5
fea : oe Le
sree: By G2. ite ee
‘ a é he ru
2 “| ay a \ Pre Nee
i . f “Vv
ri 4
‘ iu F + Yo
; : iat ’ i a 2
Whe =
tie : =“
Aly La { PAS
' r '
H =F ae y%
& THs ;
4 ? : ’ 4 igo
, ‘ t } 4 ) ;
: rie: FY a oe e) rie
"hy Pa la
i % Voom, y \s 4
is + .
1 ti hg AKG. } te j
P eat . 4 ¥ iad 4 ei iy!
rt
\
: Fs
i ,
: fi
e "
t '
;
i
ri
+ Yi ; 4
sf tae
7 ‘ f hy PA ta “rw Ae
’ , b ea) '
‘ 7 Bed bis Fr ilieae {ity ;
; ai ) bh 4 mi
bay AN ie See ue kale”, t

| a

MYCOTAXON

hr ae eS A er aenmnnnet eee erer) FO. Pe VES)
Volume LX, pp. 275-281 October-December 1996

NOTES ON CONIDIAL FUNGI. X. A NEW SPECIES OF

CERATOSPORELLA AND SOME NEW COMBINATIONS

R. F. Castafieda Ruiz', J. Guarro”, J. Cano’

' Instituto de Investigaciones Fundamentales en Agricultura Tropical "Alejandro Humboldt" (INIFAT)
Calle 1 esq. 2 Santiago de las Vegas, Ciudad Habana, Cuba. * Unitat de Microbiologia, Facultat de
Medicina i Ciéncies de la Salut, Universitat Rovira i Virgili, 43201 Reus, Tarragona, Spain

ABSTRACT

Ceratosporella compacta anam. sp. nov., found on a decaying stem of an
unidentified member of the Poaceae, is described and illustrated. It is distinguished
by its compact, dark brown to black conidia with 4 - 5 arms closely appressed.
Each arm ends in a short, robust appendage toward the apex, sometimes with a
mucilaginous tunica. Some species hitherto described in Ceratosporella are
recombined.

Key words:Cerafosporella, Acrodictys, Actinocladium, Arachnophora,
Triposporium, hyphomycetes, systematics, tropical fungi

RESUMEN

Se describe e ilustra Ceratosporella compacta anam. sp. nov., aislada
sobre restos de hojas de una especie no identificada de Poaceae. Se distingue por
sus conidios compactos, de color pardo oscuro a negro, con 4 - 5 brazos
estrechamente agrupados. De cada extremo del brazo emergen cortos y robustos
apéndices con una envoltura mucilaginosa. Algunas especies anteriormente
descritas en Ceratosporella son reclasificadas.

Palabras clave: Ceratosporella, Acrodictys, Actinocladium, Arachnophora,
Triposponum, systematics, tropical fungi

276

Ceratosporella compacta Castafieda Ruiz, Guarro et Cano anam. sp. nov.

(Fig. 1,2)
Ad fungos conidiales, hyphomycetes pertinens. Coloniae effusae, nigrae.
Mycelium partim superficiale et partim in substrato immersum, ex hyphis septatis,
ramosis, laevibus, atrobrunneis, 2 um diam compositum. Conidiophora conspicua,
mononematosa, eramosa, erecta, recta, cylindrica vel subulata, leviter inflata ad
basim, O - 2 septata, levia, nigra, 18 - 48 x 6 - 10 um. Cellulae conidiogenae
monoblasticae, terminales, interdum percurrentes, cylindricae, in conidiophoris
incorporatae. Conidia cheiroidea, compacta, acrogena, solitaria, levia, atrobrunnea
ad basim, brunnea ad apicem, ex cellula basalis et quator vel quinque ramulis
composita; cellula basalis cylindrica, truncata, nigra vel atrobrunnea, levia, 5 - 8 x 5
- 7 wm; ramuli plus minusve cylindnci ad basim, attenuati ad apicem, curvati,
adpressi, 12 - 14 septati, 65 - 70 um longi et 4 - 5 um crassi (appendice exclusa),
insidentes ad basim et apicem, sed leviter divergentes mutati in appendicibus
cellularis. Appendicibus cylindricis, inflatis et rotundatis ad apicem, 2 - 6 septatis, 15
- 46 x 2.5- 4 um, brunneis, pallide brunneis ad apicem, robustis, ex interdum tunica
mucosa, 6-12 um diam ad apicem praeditus. Teleomorphosis ignota.

Conidial fungi, hyphomycetes. Colonies effuse, black. Mycelium superficial and
immersed. Hyphae septate, branched, smooth, dark brown, 2 um diam.
Conidiophores differentiated, mononematous, simple, erect, straight, cylindrical or
subulate, slightly inflated at the base, 0 - 2 septate, smooth, black, 18 - 48 x 6 - 10
um. Conidiogenous cells monoblastic, terminal, cylindrical, integrated, sometimes
percurrent. Conidia cheiroid, acrogenous, solitary, smooth-walled, dark brown at
the base, brown at the apex; composed of a basal cell and 4 - 5 arms; basal cell
cylindrical, truncate, black or dark brown, smooth, 5 - 8 x 5 - 7 um; arms more or
less cylindrical, attenuate toward the apex, curved, appressed, 12 - 14 septate, 65 -
70 x 4 -5 um (appendage excluded), slightly divergent toward the base and the
apex, but concurrently turning progressively into cellular appendages in apical
region; appendages cylindrical, inflated and rounded at the apex, robust, 2 - 6
septate, 15 - 46 x 2.5 - 4 um, pale brown, sometimes with a slimy tunica of 6 - 12
um diam at the apex. Teleomorph unknown.

Holotype: INIFAT C94/162, on fallen decaying stem of unidentified member of the
Poaceae, Vifhales, Pinar del Rio, Cuba,11 November 1994, R. F. Castaneda.
Isotype: MUCL 39130 (Mycotheque Universite Catholique de Louvain). Cultures ex
type: INIFAT C94/162, CBS 184.95 and MUCL 39130.

277

ee,

Fig.1- Ceratosporella compacta (INIFAT C94/ 162). Conidiophores and conidia (bar

um)

278

Among the previously described species of Ceratosporella, only C. stipitata
(Goidanich) S.J.Hughes (1952), C. caliculata Lutrasti (1980); C. ponapensis
Matsushima (1981) and C. fertilis Castaneda Ruiz (1985) resemble C.compacta
superficially. However, C. stipitafa has arms up to 130 um long and 6 - 9 um thick,
without appendages; C. caliculata has 6 - 7 septate arms and a verrucose outer
wall; C. ponapensis has 4 - 7 arms up to 120 um long and lacks appendages and
C. fertilis has usually 8 - 10 arms, 33 - 60 um long, usually without appendages.

Ceratosporella deviata Subramanian (1957) has stauroconidia which strongly
resemble Tnposponum Corda, especially with T. elegans Corda (Ellis 1971), as
suggested by Sinclair et al. (1987); and the following new combination is proposed:

Triposporium deviatum (Subramanian) Castaheda Ruiz, comb. nov.
Basionym: Ceratosporella deviata Subramanian in Proceeding of the Indian
Academy of Sciences, Sect. B 46: 327 (1957).

Two species of Ceratosporella, C. longiramosa Castaneda Ruiz (1988) and C.
pulneyensis Subramanian & Bhat (1987) lack cheiroid conidia. The first has
stauroconidia with 3 divergent, distoseptate, 65 - 160 um long arms and which
would be better classified in Actinocladium Ehrenb.: Pers. The latter has stellate
conidia with two cells, each bearing 2 divergent arms, 14 x 3 - 5 um. It should be
Classified in Arachnophora Hennebert (1963). Hence the following new
combinations are proposed:

Actinocladium longiramosum (Castafieda Ruiz) Castafteda Ruiz, comb.

nov.

Basionym: Ceratosporella longiramosa Castafieda Ruiz in Fungi Cubenses
Ill, Instituto de Investigaciones Fundamentales en Agricultura
Tropical "Alejandro de Humboldt", p 2 (1988) Cuba

Arachnophora pulneyensis (Subramanian & Bhat) Castafeda Ruiz,

comb. nov.

Basionym: Ceratosporella pulneyensis Subramanian & Bhat in Kavaka 15:
50 (1987)

Ceratosporella lambdaseptata Matsush. (1971) has stauroconidia with 3-4 radial
arms and has been placed in Tnposporium by Kuthubutheen and Nawawi (1991),
and Ceratosporella goidanichii Rambelli (1958) has cylindrical conidia without arms
and has been relocated in Spondesmium by Hughes (1979). Other previously
described Ceratosporella spp. are: C. bicornis (Morgan) Hohnel (Ellis 1971), C.
novae-zelandiae S.J. Hughes (1971), C. cheiroidea Sinclair, Morgan-Jones &
Eicker (1987), C. disticha Kuthubutheen & Nawawi (1991), C. basibicellulana
Matsush. (1993), C. basicontinua Matsush. (1993) and C. flagellifera Matsush.
(1993). Acrodictys furcata Ellis (1963) has cheiroid conidia which resemble C.
novae-zelandiae and could be classified as Ceratosporella, but no new
combination is proposed.

219

We propose the following key for the identification of Ceratosporella species:

KEY TO SPECIES OF CERATOSPORELLA

aris (OrniCica: SINOOU Walled isccocit anes <. Mapanheliaed Ueto end tos tcnicacie ses 2,
1’. Conidia verrucose, with 4 arms. Arms 6-7 septate,
a) SAC, Ve Uy es TTL Sod 01 9 ae NE OM PEAR PA Lees at A oR C. caliculata

2. Conidia with 2 divergent arms forming angles
wider than 45°. Arms 5 - 7 septate, 50 - 80 x 9 - 13 um C. bicornis

2’. Conidia with no divergent ArMS ...............ccecceecceeeeeee eens 3
3. Conidia usually with more than 4 armsS..........0.0...00cceceeeeee 4
3’. Conidia usually with 4 or less ArMS.............0.ccceceeceseeeeeeee 6

4. Conidial arms 65 - 70 um long, 12 - 14 septate

PANO WEIN AD DOMOaGeS i ).8 sas meet i Ga malin 9) C. compacta
4. Conidial arms shorter than 65 [WM ..........0.0ccccceeeceeeeeeee 5
5. Conidia with 5 - 8 arms. Arms 4 - 8 septate,
Ee re OC ANTE ONG ci ta ee ain eR ie Nee SN RE oe, C. ponapensis
5’. Conidia with 8 - 12 Arms. Arms 7 - 14 septate,
COON NORA eee: staples iecct Sam Ovook eee Mn tau thar C. fertilis
6. Conidia with apical appendages ............0...c cece if
6’. Conidia without appendages ..............0..0. cece eeteeeeeeees 8
7. Conidia with septate appendages;
UAMIS WTS) 0 SEP ed oie ei teh he ee ascot oe cl arabe oa cee eae tees C. flagelligera
7’. Conidia with non-septate appendages;
alinisiwithy'O — 10 Septet). & Nee Aceh tian |. 8 Sead tv adlaes C. disticha
8. Conidia with the basal cell darker than the rest ................ C. cheiroidea
BZ OP ICEIIOL AS QOOVE sos cccisha vetintie seed svn ue dauod nebant aeeet sc 9
9. Conidia usually with 3 - 4 arms, 6 - 8 um wide..................05. C. stipitata
Die COnicia: always: WIthl.2 ALINS .F.25. oe cicteees Seeeel cic tc Vaevettent oes 10

10. Basal part of the conidia 1-septate;
conidial arms 2 - 6 septate ............0 ee eeteeeereeteees C. basibicellulana
10’. Basal part of the conidia aseptate .......... eee 11

11. Conidial arms emerging symmetrically from

tne pasar ce). |= oO SOPtale ne ees nai uc o oec ne aetsacte C. basicontinua
11’. Conidial arms emerging asymmetrically from

the basal cell (one of them in lateral position), 1 - 3 septate,

slightly constricted at the septa... cee eeees C. novae-zelandiae

280

Fig.2- Ceratosporella compacta (INIFAT C94/162). a,b, conidiophores and conidium, ¢,d,
cellular appendages (bar: a = 40 um, b = 25 ym, c,d, = 10 ym).

ACKNOWLEDGEMENTS

We gratefully acknowledge Dr. E. Descals (Inst. Mediterrani d’ Estudis Avangats de
les Balears, C.S.I.C.) for serving as pre-submission reviewers, The Cuban Ministry
of Agriculture for facilities during this work. This work was supported the
“Fundacion Ciencia i Salut’, Reus, Spain.

281

REFERENCES

Castaneda Ruiz, R.F. (1985). Deuteromycotina de Cuba. Hyphomycetes. II.
Instituto de Investigaciones Fundamentales en Agricultura Tropical
"Alejandro de Humboldt", Cuba, 23 pp.

Castaneda Ruiz, R.F. (1988). Fungi cubenses Ill. Instituto de Investigaciones
Fundamentales en Agricultura Tropical "Alejandro de Humboldt", Cuba, 54

pp.
Ellis, M. B. (1963). Dematiaceous Hyphomycetes. V. Mycol. Pap. 93: 1-33.

- Ellis. M. B. (1971). Dematiaceous Hyphomycetes. Commonth. Mycol. Inst., Kew,

608 pp.

Hennebert, G.L. (1963). Un hyphomycete nouveau, Arachnophora fagicola gen.
nov., sp. nov. Can. J. Bot. 41: 1165-1169.

Hughes, S.J. (1952). Speira stipitata. Trans. Brit. Mycol. Soc. 35: 243-247.

Hughes, S.J. (1971). New Zealand Fungi 16. Brachydesmiella, Ceratosporella.
N. Z. J. Bot. 9: 351-354.

Hughes, S.J. (1979). Relocation of species of Endophragmia auct. with notes on
relevant generic names. WN. Z. J. Bot. 17: 139-188.

Lustrati, L. (1980). Ceratosporella caliculata, sp. nov. nuova specie di ifale
demaziaceo. Mic. Ital. 3: 11-14.

Kuthubutheen, A. J. & Nawawi A. (1991). A new species of Ceratosporella
and Tnposponum lambdaseptatum (Matsush.) comb. nov. from Malaysia.
Mycol. Res. 95: 158-162.

Matsushima, T. (1971). Microfungi of the Solomon Islands and Papua-New
Guinea, Kobe. Matsushima.

Matsushima, T. (1981). Matsushima Mycological Memoirs No.2, Kobe.
Matsushima.

Matsushima, T. (1993). Matsushima Mycological Memoirs No.7, Kobe.
Matsushima.

Rambelli, A. (1958). Micromiceti della Foresta di Campigna. 11° Contributo. Atti
Accad. Scienze Ist. Bologna, cl. Sci. Fis., ser.Xl, §:1-15.

Sinclair, R. C., Eicker, A. & Morgan-Jones G. (1987). Notes on :
Hyphomycetes. LVI. Ceratosporella cheiroidea, a new species. Mycotaxon
30: 351-355.

Subramanian, C. V. (1957). Hyphomycetes IV. Proc. Indian Acad. Sci., sect.B,
46: 324-335.

Subramanian, C. V. & Bhat, D.J. (1987). Hyphomycetes from South India |.
Some new taxa. Kavaka, 15: 41-74.

yu 7 hy
my P) ) a '
4 A fi 1
s , yy
(
fh:
, fa
e ;
As a |
\* My ays.
: en
: Te aa
¥, hea ‘ \
; 4
Evo pgs
v SOW ALY wan we
t ee! f ‘ Ct
, a a ya
; ok iw ae
y ew

ihr me 4

y
yn’ at
wh ‘4,
eK sgh sp bi
Ades hanes

rstiay 4h a a \Giud tes a Guat a
at ade elites en Niky iia

inh es Mae da bits
{2 ‘ iheitay "te * A te ;
Tae be i) NPA TO ah: mn

\ * At ak) Pr kal

, ‘ j
Ay ight i}
J i cee vat "

MYCOTAXON

a ee
Volume LX, pp. 283-290 October-December 1996

SOME AIRBORNE CONIDIAL FUNGI FROM CUBA

R.F. Castafieda Ruiz’, D.E. Fabré*, M.P. Parra®, M. Pérez’, J. Guarro®

and J. Cano®

‘Instituto de Investigaciones Fundamentales en Agricultura Tropical "Alejandro de Humboldt" (INIFAT) Calle 1,
esq. 2 Santiago de las Vegas. Ciudad Habana. Cuba. "Hospital Pediatrico Docente "Juan Manuel Marquez” Avenida
31. Marianao, Ciudad Habana, Cuba. *Unitat de Microbiologia, Facultat de Medicina i Ciéncies de la Salut,
Universitat Rovira i Virgili. 43201 Reus, Espafia

ABSTRACT

Zeloaspensponum hyphopodioides anam. gen. et sp. nov. is described and
illustrated from conidia in the atmosphere of Cuba. This fungus is distinguished by
the short and clearly differentiated hyphopodia arising from vegetative hyphae and
0-2 septate, verrucose conidia arising from sympodial conidiogenous cells. Seven
other species of hyphomycetes are cited from the same source.

Key words: Aeromycology, Zeloasperisporium hyphopodioides, hyphomycetes,
Cuba

RESUMEN

Se describe e ilustra Zeloasperisporium hyphopodioides anam. gen. et sp. nov.
aislado de conidios en la atmdésfera de Cuba. Dicho hongo se caracteriza por la
formaci6n de hifopodios cortos y bien diferenciados a partir de las hifas
vegetativas y por poseer conidios 0-2 septados y verrucosos formandose sobre
células conidi6genas simpodiales. Otras siete especies de hongos conidiales
aislados también del aire de Cuba son citadas.

Palabras clave: Aeromycology, | Zeloaspensponum —_ hyphopodioides,
hyphomycetes, Cuba

284

INTRODUCTION

During December 1992, January-February 1993, and October-November 1994, a
survey of airborne fungi possibly related to asthmatic and allergic diseases was
conducted in Cuba. Air samples were collected from 3 localities in Havana City
Province (Marianao, Playa and Cerro). In this paper only new and uncommon
conidial fungi are treated.

MATERIAL AND METHODS

Colonies were obtained on gravity exposed plates. Three plates were exposed
daily and simoultaneously for 15 minutes in and out of three different homes in
three different areas of Havana City Province. The culture medium used was SNA
(Nirenberg, 1976), with 300 ppm chloramphenicol. The plates were incubated at 25
+ 2 °C for four days. Some strains were subcultured on leaf litter during four weeks,
according to Nag Raj (1993). A more complete description of the method and
analysis of the data will be given in a later paper.

TAXONOMIC PART

Zeloasperisporium Castafeda Ruiz anam. gen. nov.
Etym.: Zelo-, from the Greek, meaning emulation; -aspensponum, a genus name in
the Fungi.

Hyphomycetes pertinens. Coloniae effusae, olivaceae usque brunneae. Mycelium
plerumque superficiale. Hyphae septatae, brunneae usque olivaceae. Hyphopodia
brevissima, "appressona" similia, tuberculata usque lobata ad apicem, saepissime
adsunt. Conidiophora conspicua, mononematosa, erecta, septata, brunnea usque
Olivacea, eramosa. Cellulae conidiogenae polyblasticae, sympodiales, integratae,
cicatncatae cum cicatricibus conspicuis praeditae. Conidia obclavata vel clavata,
fusiformia usque cylindnca, cicatricata, septata, levia vel verrucosa, olivacea usque
brunnea.

Species typica: Zeloasperisporium hyphopodioides anam. sp. nov.

Hyphomycetes. Colonies on natural substrate effuse, olivaceous to brown.
Mycelium mostly superficial. Hyphae septate, brown to olivaceous. Hyphopodia
short, appressorium-like, inflated, slightly warted to lobed at the apex, present.
Conidiophores differentiated, mononematous, erect, septate, brown to olivaceous,
unbranched. Conidiogenous cells polyblastic, sympodial, terminal, integrated, with
evident dark detachment scars. Conidia obclavate, clavate, fusiform, or cylindrical,
septate, verrucose or smooth, olivaceous to brown, scarred.

285

Zeloaspensponum shows some resemblance with Passalora Fries, Asperisporium
Maublanc, and Fusicladium Bonorden (Ellis 1971, 1976; v. Arx 1983,1987). All
have blastic conidiogenesis and sympodial, terminal, cicatrized conidiogenous
cells; and obclavate, cylindrical to fusiform or obovoid conidia. However, vegetative
hyphae in Passalora are immersed in the host tissue and form
pseudoparenchymatous stromata, whereas in Asperisponum and Fusicladium the
hyphae form well-developed, erumpent stroma. Hyphopodia are only present in
Zeloaspensponum

Zeloasperisporium hyphopodioides Castafieda anam. sp. nov. Figs. 1,2

Ad hyphomycetes pertinens. Coloniae in foliis putridis sterilifactis Bucidae palustris,
effusae, amphigenae, olivaceae. Mycelium plerumque superficiale. Hyphae
septatae, ramosae, plerumque verrucosae, interdum leves, brunneae usque
Olivaceae, 3.5 mm diam, Hyphopodia brevissima “appressoria" similia, cylindrica,
2.5-45x 1 - 1.5 um, inflata, leviter tuberculata usque lobata ad apicem, pallide
olivacea et obscura ad apicem, 2.5 - 4 x 7 - 1.5 mum, interdum numerosa,
saepissimae adsunt. Conidiophora conspicua, mononematosa, subulata, erecta,
recta, O - 1 septata, levia vel leviter verrucosa, brunnea vel olivacea, eramosa,
usque 40 um alta et 3.5 - 4.5 um crassa ad basim. Cellulae conidiogenae
polyblasticae, sympodiales, integratae, cicatricatae cum cicatncibus obscuns,
conspicuis praeditae. Conidia obclavata, conico-truncata, ad basim cicatrcata,
obtusa ad apicem, 0-2 (1) septata, constrcta ad septum, verrucosa, olivacea vel
brunneo-olivacea, sicca, 16 - 25 x 3 - 4 um.Teleomorphosis ignota.

Hyphomycetes. Colonies on sterilized leaves of Bucida palustris effuse,
amphigenous, olivaceous. Mycelium mostly superficial. Hyphae septate, branched,
verrucose, sometimes smooth, brown to olive, 2 - 3 mm diam. Hyphopodia short,
cylindrical, inflated, slightly warted to lobed at the apex, pale olive, darker at the
end, 2.5 - 4 x 1 - 1.5 um, sometimes numerous. Conidiophores differentiated,
mononematous, subulate, erect, straight, O-1 septate, smooth-walled or slightly
verrucose, brown to olive, unbranched, up to 40 um tall, 3.5 - 4.5 um wide at the
base. Conidiogenous cells polyblastic, sympodial, terminal, integrated, with
conspicuous dark scars. Conidia obclavate, obconical truncate at the base,
scarred, obtuse at the apex, 0-2 (1) septate, constricted at the septa, verrucose,
olive or brown-olivaceous, dry, 16 - 25 x 3 - 4 um. Teleomorph unknown.

Type specimen: On sterilized leaves of Bucida palustris, C. Havana. Cuba, R.F.
Castaneda, 4 October. 1994. Holotype: INIFAT C94/114, in INIFAT Herbarium,
Havana, Cuba. Isotype: CBS 257.95. Living cultures ex type: INIFAT C94/114,
CBS 218.95, MUCL 39155 and IMI 367520.

Fig. 1.- Zeloasperisporium hyphopodioides (C94/114). a-c, hyphopodia (a, on natural
substrate; b,c, on SNA). d-g conidiophores, conidiogenous cells and conidia.
Conidophore and conidia (f, on natural substrate; g, on SNA). h, conidia (a,d,fh,

bar = 10 pm; b,c,e,g, bar = 5 pum).

287

Fig. 2.- Zeloasperisporium hyphopodioides (C94/114). Conidiophores, conidiogenous cells
and conidia (a, on SNA; b, on natural substrate) (bar = 10 pum)

288

OTHER AIRBORNE CONIDIAL FUNGI COLLECTED.

Arxiella terrestris Papendorf., Trans. Br. Mycol. Soc. 50: 73 (1967)
Specimen examined: INIFAT C93/112, from air, Playa. C. Havana, Cuba, 10. Aug.
1994, R.F. Castafeda.

Botryotrichum peruvianum Matsushima, Icon. microf. a Matsushima lect.,
Kobe: 17 (1975)
Specimen examined: INIFAT C93/233 from air, C. Havana, Cuba, 12, Feb. 1993,
R.F. Castaneda.

Curvularia tuberculata Jain, Trans. Br. Mycol. Soc. 45: 539 (1962) Fig.3 a,b
Specimen examined: INIFAT C94/107 from air, C. Havana, Cuba, 4, Oct. 1994,
R.F. Castaheda.

Curvularia verrucosa Sivanesan, Mycol. Res. 96: 485 (1992)
Specimen examined: INIFAT C93/234 from air, C. Havana, Cuba, 12, Feb. 1993,
R.F. Castaneda.

Dactylaria kumamotoensis Matsushima, Matsush. Mycol. Mem. 2: 5 (1981).

Fig.3 c,d
Specimen examined: INIFAT C94/128, from air, C. Havana, Cuba, 4, Oct. 1994,
R.F. Castaneda.

Dwalomyces taiwanensis Matsushima, Matsush. Mycol. Mem. 5: 14 (1987)
Specimen examined: INIFAT C93/57, from air, Playa, C. Havana, Cuba. 9,
Feb. 1993. R.F. Castaneda.

Tritirachium oryzae (Vincens) de Hoog, Stud. Mycol. 1: 22 (1972)
Specimen examined: INIFAT C93/97, from air, Ciudad Habana, Cuba, 9,
Feb. 1993. R.F. Castafeda.

ACKNOWLEDGEMENTS

We gratefully acknowledge Dr. E. Descals (Inst. Estudis Avancats de les Balears,
C.S.I.C.) for serving as pre-submission reviewers, The Cuban Ministry of
Agriculture for facilities during this work. This work was supported by the
“Fundacion Ciencia i Salut’, Reus, Spain.

Fig.3.- Curvularia tuberculata (INIFAT C94/107). a,b (a, bar = 25 um; b, bar = 10 um).
Dactylaria kumamotoensis (INIFAT C94/128). c,d (bar = 10 um)

290

REFERENCES

Arx, J.A.von (1983). Mycosphaerella and its anamorphs. Proc. K. Ned. Akad.
Wet.,c, 86: 15-54.

Arx, J.A.von (1987). Plant pathogenic fungi. Beihefte zur Nova Hedwigia, 87: 1-
188

Ellis, M.B. (1971). Dematiaceous hyphomycetes. Commth. Mycol. Inst. Kew. 608

Pp.

Ellis, M.B. (1976). More Dematiaceous hyphomycetes. Commth. Mycol. Inst.
Kew. 507 pp. .

Nag Raj,T.R.(1993). Coelomycetous anamorphs with appendage-bearing
conidia. Mycologue Publications, Waterloo, 1101 pp.

Nirenberg,H.1.(1976).Untersuchungen Uber die morphologische und biologische
Differenzierung in der Fusanum-Sektion Liseola. Mitteil. Biol.
Bundesanst. Land-Forstw, Berlin-Dahlem 169,117 pp.

MYCOTAXON

EGYPTIAN UREDINALES. I. RUSTS ON WILD PLANTS
FROM THE NILE DELTA.

ZAKARIA A. BAKA! AND HALVOR B. GJ4RUMZ2

1. Department of Botany, Faculty of Science, University of Mansoura, Egypt.
2. Planteforsk, Plant Protection Centre, Fellesbygget, N-1432 AS, Norway.

ABSTRACT

Twenty-three rust species on moncotyledonous and dicotyledonous
wild plants were recorded in the Nile Delta, Egypt, from the period of
1991-1994. One Melampsora species, eighteen Puccinia species, and
four Uromyces species were identified. Eight Puccinia species are
recorded as new to Egypt, viz. Puccinia striiformis on Lolium perenne,
P. polypogonis on Polypogon monspeliensis, P. fragosoana and P.
imperatae on Imperata cylindrica, P. cynodontis on Cynodon dactylon,
P. cyperi-tegetiformis on Cyperus rotundus, P. isiacae on Phragmites
australis and P. lagenophorae on Senecio glaucus and S. aegyptius.

El-Helaly et al. (1966) published the only report about the names
of some rust species and their hosts in Egypt up to 1965 without detailed
descriptions of the fungi.

Melampsora euphorbiae (Schub.) Cast., Obs. Pl. Acotyl. 2:18
(1843).

Syn. M. helioscopiae Wint. in Rabh. Krypt.-Fl., 2, 1, 1:240 (1882); M.
euphorbiae-gerardiana W. Muller.

On Euphorbia peplus L., E. helioscopia L., E. heterophylla L.
(Euphorbiaceae), II+IIT.

Localities: Univ. Mansoura farm, Mansoura-Zagazig Rd.

Uredinia amphigenous, hypophyllous with capitate, thick-walled,
hyaline paraphyses. Urediniospores globoid or ellipsoid, 18-22 x 12-20
um, with paraphyses intermixed, wall colorless, 2-3 um thick, finely
echinulate, pores invisible. Telia amphigenous and occasionally
caulicolous, dark brown, subepidermal., Teliospores prismatic, 30-44
x 7-13 um, wall yellowish brown 1-1.5 um thick.

292

Many Melampsora species have been described on Euphorbia
spp., but according to Jorstad (1958) "it seems impossible to draw
definite lines of demarcation between the various species". M.
euphorbiae s. lat. is widespread nearly all over the world. The first
record of this rust in Egypt was given by v. Thumen (1979) and
Roumeguere (1881) on Euphorbia sp. Later it has been reported on E.
arguta Soland., E. cornuta Pers., E. peploides Gaven and E. peplus L.
(cfr. El-Helaly et al. 1966). E. helioscopia and E. heterophylla are new
hosts for the rust in Egypt, but they have been reported as such from
other countries, including Africa. Castellani & Ciferri (1937) reported
it on E. monticola Hochst. in Eritrea and later Castellani & Ciferri
(1950) on E. schimperiana Scheele from Sidamo. Bisby & Wiehe
(1953) and Gjzrum (1977) have reported it on E. depauperata A.
Rich. in Ethiopia and Malawi, respectively. Stewart & Yirgou (1967)
and Gjzrum (1985) have reported it on E. heterophylla from
Zimbabwi and Uganda, respectively.

Puccinia cancellata Sacc. & Roum., Rev. Myc. 9:26 (1881).

Syn. Uredo cancellata Dur. & Mont., Fl. Alger. 1:314 (1846).

On Juncus acutus L. (Juncaceae), II+III.

Localities: Baltim, Rashid, Mansoura-El-Mahala Rd.

Uredinia large, elongate, surrounding the stem, dehiscing with a
longitudinal slit, chestnut-brown. Urediniospores ellipsoid, pale brown,
32-39 x 19-26 um, wall 2.5-4 um, echinulate, 3 equatorial germ pores.
Telia mixed with uredinia. Teliospores oblong, pale brown, constricted
at septum, 34-47 x 20-27 um; wall 3-4 um thick at sides, 6-7 um at
apex, smooth; pedicels hyaline, slender, deciduous, up to 45 um.

J. acutus is the main host for this rust, known from many countries
around the Mediterranean Sea, going east to Iran. It ts also reported
from the Canary Islands, Madeira, Azores and Great Britain. Reichert
(1921) reported P. rimosa Wint. (= P. cyrnaea Maire) on J. acutus, J.
maritimus Lam. and J. subulatus Forsk. (cfr. Melchers 1931 and EI-
Helaly et al. 1966). As J. maritimus is the main host of P. cyrnaea this
record might be doubtful.

293

Puccinia carthami Corda, Ic. Fung. 4:15 (1840).

Syn. Bullaria carthami (Corda) Arth. & Mains, N. Am. Fl. 7:512
(1922).

On Carthamus tinctorius L. (Asteraceae), II.

Localities: Kalabshu, Baltim.

Uredinia amphigenous, scattered, brown. Urediniospores globose, 22-26
x 20-24 um; wall chestnut-brown, 1.5-2 um thick, echinulate, 2-3
equatorial germ pores.

C. tinctorius is the main host for this rust in N. America, Europe,
Australia and Africa. It is also reported on other Carthamus spp.,
especially in Asia, but also in Portugal, Spain and Morocco. In infection
experiments Parmelee (1960) got very week infections on Centaurea
cyanus L. From Egypt it was first reported by Reichert (1921) and
Snell (1923). Reichert’s record is also mentioned by Melchers (1931)
and El-Helaly et al. (1966).

Cummins (1977) made this rust species synonymous with P.
calcitrapae DC. var. centaureae (DC.) Cumm.

Puccinia cenchri Diet. & Holw. var. africana Cumm., Torrey Bot.
Club Bull. 79:217 (1952).

Syn. Uredo cenchricola P. Henn., Mus. Congo Anal. 2:223 (1908).

On Cenchrus biflorus Roxb. (Poaceae), II.

Localities: Damietta-Port Said Rd., Univ. Mansoura farm, Ismailia.
Uredinia amphigenous, cinnamon-brown. Urediniospores ellipsoid, 29-
38 x 22-30 um; wall cinnamon-brown, echinulate, 2-3 um thick, 4-5
equatorial germ pores.

P. cenchri var. africana is reported from many African countries,
often as Uredo cenchricola P. Henn. It is new to the rust flora of Egypt.

Puccinia cynodontis Lacroix ex Desm., Pl. Crypt. Ser. II, No. 655.
(1859). |

Syn. Uredo eleusine-indicae Saw., J. Taihoku Soc. Agr. For. 7:41
(1943).

On Cynodon dactylon (Pers.) L. (Poaceae), II + III.

Localities: Ismailia, Tanta, Baltim.

294

Uredinia on abaxial surface, cinnamon-brown. Urediniospores globoid,
21-27 x 20-23 um, wall 2-3 um thick, light brown, verrucose, 2-3
equatorial germ pores. Telia amphigenous, hypophyllous, dark brown,
elongated. Teliospores ellipsoid, with papillate tip, 29-52 x17-21 um,
the wall is 1.8-2.4 um thick at the sides, 4.5-11 um at apex, pedicels
light brown, thin-walled.

P. cynodontis, a heterocious rust species with aecia (Aecidium
plantaginis Ces.) on hosts belonging to many plant families, is
circumglobal in temperate and warmer areas. It is widespread,
especially in Africa, but this the first record of it in Egypt. The aecial
stage has been found on Adonis sp. in Tunisia and on Plantago
coronopus L. in Algeria.

Puccinia cyperi-tegetiformis Kern, Mycologia 11:134 (1919).
Syn. Uredo cyperi-tegetiformis P.Henn., Bot. Jahrb. 34:593 (1905).

On Cyperus rotundus LL. (Cyperaceae), II.

Locality: Univ. Mansoura farm.

Uredinia mostly on abaxial surface, elongated, cinnamon-brown.
Urediniospores pale brown, 20-26 x 16-19 um., wall dark brown,
echinulate, 1.5-3.5 um thick, 2 equatorial germ pores.

This rust species is reported on many Cyperus spp. scattered all over
Africa, and it is also known in Japan (type locality) and in the Americas.
The rust is new to the flora of Egypt, but C. rotundus has previously
been reported as a host in Uganda and Chad (Gjerum 1990).

Puccinia fragosoana Beltran, Mem. Roy. Soc. Espan. Hist. Nat.
50:249 (1921).

Syn. Uredo schizachyrii Doidge, Bothalia 2:508 (1928).

On Imperata cylindrica (L.) Raeuschel. (Poaceae), II.

Localities: Ras El-Bar, Kafr Saad, Kafr El-Batikh, Port Said.

Uredinia amphigenous, dark brown, capitate. colorless paraphyses with
thin wall and 8-15 um thick at the head. Urediniospores golden to
brown, obovoid, 31-44 x 20-28 um, wall 1.5-2.5 um thick, 3.5-7.5 um
at apex, echinulate, 4-5 equatorial germ pores.

295

P. fragosoana, widespread in Africa, and also reported from
Palestine, is new to the rust flora of Egypt. It is easily separated from P.
imperatae Poirault by the presence of uredinial paraphyses. Other
paraphysate rust species on the same host in Africa differs from P.
fragosoana having urediniospores not thickened apically.

Puccinia graminis Pers., Syn. Meth. Fung. p. 228 (1801) spp.
graminis

Syn. P. albigensis Mayor, Rev. Mycol. 22: 279 (1957).

On Cynodon dactylon (L.) Pers. (Poaceae), II.

Localities: Kafr Saad, Ras El-Bar, Kafr El-Batikh-Damietta road.
Uredinia amphigenous, cinnamon-brown. Urediniospores orange 23-39
x 15-24 um, ellipsoid, the wall is 1.5-2 um thick, echinulate, 3-5
equatorial germ pores.

P. graminis ssp. graminis, the black stem rust, is circumglobal,
occurring on many genera of Poaceae. In Egypt it has been reported on
Avena sativa L., Hordeum vulgare Lam. and Triticum spp. (Melchers
1931). C. dactylon is a new host for this rust in Egypt. The aecial stage
on Berberis spp. (Aecidium berberidis Pers.) has been reported from
Morocco and Algeria.

The other subspecies, ssp. graminicola Urban, differs from the ssp.
graminis having shorter urediniospores.

Puccinia imperatae Poirault, Assoc. Nat. Nice Bull. 1:105 (1913).
Syn. Uredo imperatae Magn., Zool.-Bot. Ges. Wien Verhandl. 50:439
(1900),

On Imperata cylindrica (L.) Raeuschel. (Poaceae), II.

Localities: Ras El-Bar, Kafr El-Batikh, Port Said.

Uredinia amphigenous, light yellow. Urediniospores hyaline, globoid,
22-30 x 20-26 um, wall yellowish, 1-2 wm thick, 3.5-7 um at apex,
echinulate, 4 equatorial germ pores.

P. imperatae is a Mediterranean rust species, described from France,
but it is also reported from S. Africa. It is new to the rust flora of

Egypt.

296

Puccinia isiacae Wint. in O. Kuntze Plantae orient.-ross. p. 127
(1887).

On Phragmites australis (Cav.) Trin. (Poaceae), II+III.

Localities: Kafr El-Batikh-New Damietta Rd., Kafr Saad El-Balad.
Uredinia amphigenous, scattered on leaves in large groups.
Urediniospores ellipsoid, 21-32 x 19-22 um; wall cinnamon-brown,
echinulate, 3-4 um thick, 3 equatorial pores. Telia amphigenous,
chocolate-brown. Teliospores brown, 35-54 x 20-30 um; wall 3-4 um
thick at sides, 5-7 um at apex; pedicels thick-walled up to 170 um long.

P. isiacae was first reported from Egypt by v. Thimen (1879) on
Arundo isiacae Del. (= Phragmidium australis (Cav.) Trin.). Later it
has been listed by Reichert (1921), Melchers (1931) and El-Helaly et
al. (1966). In Africa this rust is reported from Morocco and Tunisia. It
is reported from many countries in S. Europe and Asia.

The aecial stage occurs on hosts belonging to about 20 plant families
(Cummins 1971), but this stage is not reported from Egypt.

Puccinia lagenophorae Cooke, Grevillea 13: 6 (1884).

Syn. Puccinia terrieriana E. Mayor, Ber. Schweiz. Bot. Ges. 72:

266 (1962).

On Senecio glaucus L. and S. aegyptius L. (Asteraceae), I+III.
Localities: Baltim, Rashid, Kafr Saad El-Balad, Ras-El-Bar.

Aecia amphigenous, on stems and leaves, orange. Aeciospores
polygonal, 12 x 17 um. Wall hyaline, finely verrucose, 0.8 um thick.
Telia pulvinate, dark, sparsed on the stems. Teliospores clavate to
ellipsoid, 30-40 x 12-20 um. Wall smooth, 1.5-2 um thick at sides and
5.5-8.5 um at apex; pedicels persistent, hyaline, 23-42 x 3.5-6 um.
Some telia have up to 16% 1-celled mesospores, 18-32 x 11-16 um.

P. lagenophorae is described from Australia where it lives on many
asteraceous genera. It seems rare in Africa, but since 1960 it has spread
in South and Central Europe, and also in Norway, mainly on Senecio
vulgaris L. It is also reported from Argentina. There is no previous
report on this rust from Egypt. S. aegyptius is a new host for this rust
while S. glaucus (as ssp. coronopifolius (Maire) Alexander) has been
reported as a host from the Canary Islands (Gjzrum 1987).

297

Puccinia magnusiana Koern., Hedwigia 15:179 (1876).

Syn. Puccinia alnetorum Gaeum., Hedwigia 80:139 (1941).

On Arundo donax L. (Poaceae), II+III.

Localities: Kafr Saad El-Balad, Damietta-Ezbet El-Borg Rd.

Uredinia amphigenous, yellowish brown, with capitate, hyaline
paraphyses, 35-60 um long; wall 1.5-3.5 um at sides, 4-6 um at apex.
Urediniospores ellipsoid or oval, 22-39 x 14-19 um; wall yellowish
brown, 1.5-2.5 um thick, echinulate, 7-10 scattered germ pores. Telia
amphigenous, dark brown. Teliospores oblong to ellipsoid, 37-57 x 15-
28 um; wall chestnut-brown, smooth, 1.5-3 um thick at sides, 6-12 um
at apex; pedicels brownish, persistent, up to 60 um long.

P. magnusiana is a circumglobal rust, living on Arundo and
Phragmites. A. donax is a new host for this rust in Egypt, but it is
previously reported on Phragmites communis Trin. var. isiaca (Del.)
Cosson (= P. australis (Cav.) Trin.) (Reichert 1921). It has been
reported on A. donax in Europe and Asia.

Puccinia menthae Pers., Syn. Meth. Fung. p. 227 (1801)

On Mentha longifolia L. (Lamiaceae), II.

Localities: Rashid, Baltim, Kalabshu.

Uredinia mostly hypophyllous, irregularly scattered or occasionally in
concentric groups. Urediniospores ellipsoid, 21-27 x 16-19 um. The
wall 1.5-2.5 um thick, finely echinulate, 3 equatorial germ pores.

P. menthae is circumglobal, living on many lamiaceous host genera. In
Africa it has been reported from many countries from the
Mediterranean through East Africa to S. Africa (Gjzrum 1977). EI-
Helaly et al. (1966) reported it in Egypt on Mentha x piperita L. as did
Melchers (1931). Also, Hennings (1891,1893) and Jgrstad (1956)
have reported it on Mentha longifolia from Eritrea. Gjerum (1977)
have reported it on Satureja paradoxa (Vatke) Engl. from Ethiopia.

Puccinia polygoni-amphibii Pers., Syn. Meth. Fung. p. 227 (1801)
var. polygoni-amphibii.

Syn. Aecidium sanguinolentum Lindr., Bot. Notiser: 241 (1900).

On Polygonum salicifolium Brouss. ex Willd. (Polygonaceae), II.

298

Locality: Damietta-Kafr El-Batikh Rd.

Uredinia amphigenous, mostly hypophyllous, rounded, brownish.
Urediniospores ellipsoid, 18-32 x 16-21 um; wall cinnamon-brown,
echinulate, 1-1.5 um thick, 2-3 supra-equatorial germ pores.

This rust taxon is circumglobal, mainly on species of Polygonum, but
it occurs also on the related genus Oxygonum.. In Africa it is reported
scattered from Morocco, Algeria and Libya, but we have seen no record
from Egypt. The aecial stage (Aecidium sanguinolentum Lindr.) has
been reported by Maire (1905) on Geranium atlanticum Doiss from
Algeria.

Puccinia polypogonis Speg., An. Mus. Nas. B. Aires 19:300
(1909).

Syn. Uredo polypogonis Speg., An. Mus. Nas. B. Aires 6:240 (1899).
On Polypogon monspeliensis (L.) Desf. (Poaceae), II.

Localities: Alexandria, Faraskur.

Uredinia on abaxial leaf surface, cinnamon-brown. Urediniospores
obovoid, golden, 23-29 x 22-25 um. The spore wall echinulate, 1.5-2
um thick, 5-7 scattered germ pores with conspicuous caps.

P. polypogonis, described from Argentina, occurs on Polypogon ssp.
in S. America and S. Africa. It is new to the rust flora of Egypt.

Puccinia recondita Roberge ex Desmazieres, Bull. Soc. Bot. France
4:798 (1857).

Syn. Puccinia dasypyri Guyot & Malen., Trav. Inst. Sci. Cherif Ser.
Bot. 28:62 (1963).

On Lolium perenne L. (Poaceae), II.

Localities: Kafr Saad El-Balad, Ismailia.

Uredinia amphigenous, cinnamon-brown. Urediniospores broadly
obovoid, 23-35 x 19-27 um; wall 1-2 um thick, yellowish brown,
echinulate, 6-9 scattered germ pores.

P. recondita, a circumglobal rust species, most common in temperate
areas, is treated here as an aggregate species, occurring on a large
number of host genera of Poaceae. As P. dispersa Eriks. & P. Henn. it

299

has been reported from Egypt on L. temulentum L. by Reichert
(1921). L. perenne is a new host for this rust in Egypt.

Puccinia scirpi DC., Fl. Fr. 2:223 (1805).

On Bolboschoenus maritimus (L.) Palla (syn. Scirpus maritimus L.)
(Cyperaceae), II.

Localities: Talkha-Belqas Rd., Damietta-Port Said Rd.

Uredinia scattered or in rows, oblong, brown. Uredinospores ovoid to
sub-globoid, flattened on one side, 20-30 x 11-25 um; wall pale brown,
echinulate, 2-3 um thick, 2 equatorial germ pores.

Reichert (1921) reported this rust on Scirpus sp. while Melchers
(1931) and El-Helaly et al. (1966) reported it on the host mentioned
above in Egypt. From Morocco and Tunisia it is reported on
Schoenoplectus lacustris (L.) Palla (syn. Scirpus lacustris L.).

The aecial stage occur on species of Limnanthemum and
Nymphoides, but to our knowledge it is not reported from Africa.

Puccinia striiformis Westend., Bull. Roy. Acad. Belg., Cl. Sci.
21:235 (1854) var. striiformis.

Syn.Uredo glumarum J. K. Schmidt, Allgem. Oekon.-tech., Fl. 1:27
(1827).

Puccinia stapfiolae Mundk. & Thirum., Imp. Mycol. Inst. Kew Mycol.
Papers 16:14 (1946).

On Lolium perenne L. (Poaceae), II+III.

Localities: Univ. Mansoura farm, Zagazig.

Uredinia amphigenous, in linear series in chlorotic streaks, orange, with
saccate paraphyses, 12.5-24 um diam., wall colorless, 0.5 um thick.
Uredinispores broadly ellipsoid, 22-31 x 17-25 um; wall pale yellowish,
1.5-2 um thick, echinulate, 9-13 scattered germ pores. Telia on abaxial
leaf surface, with brown paraphyses. Teliospores oblong, 35-63 x 14-23
um.; wall deep golden brown, smooth, 1.5-2 um at sides and 5-9 um
apically; pedicels colorless to brownish, thin-walled, less than 18 um
long.

P. striiformis var. striiformis, the stripe rust, is circumglobal,
especially in the northern hemisphere, where it lives on many species of

300

Poaceae. As P. glumarum it has been reported from Egypt on Hordeum
vulgare L. and Triticum spp. (Melchers 1931).

Another variety, var. dactylidis Manners on Dactylis glomerata L.,
has smaller spores. So far it is not reported from Africa.

Puccinia tuyutensis Speg., Fungi Argent. 4:25 (1881).

Syn. P. cressae Lag. in Bol. Soc. Brot., p.131 (1889); Aecidium cressae
DC., Fl. Fr. 6:89 (1815).

On Cressa cretica L. (Convolvulaceae), I+II+III.

Localities: Baltim, Kalabshu.

Aecia hypophyllous on stems and leaves, orange. Aecidiospores globoid,
hyaline, 20-26 x 19-25 um, wall thin 1-2 um, verrucose. Uredinia
hypophyllous, cinnamon. Urediniospores ellipsoid, 23-30 x 23-25 um,
wall 2-3 um thick, echinulate, 2-3 equatorial germ pores. Telia
hypophyllous, dark brown, scattered. Teleutospores brown, 41-45 x 23-
27 um, wall 3-4 um thick and 5.5-7.5 um at apex, pedicels fragile,
hyaline.

This rust is widespread in Asia and the Americas, and also in
Mediterranean countries in Africa and as well as in Europe. Reichert
(1921) and El-Helaly et al. (1966) reported it as P. cressae on the host
mentioned above from many localities in Egypt.

Uromyces linearis Berk. & Br., J. Linn. Soc. London 14:92 (1875).
On Panicum coloratum L. (Poaceae), II+ITI. i

Locality: Univ. Mansoura farm.

Uredinia epiphyllous, cinnamon-brown, oval to elongated.
Urediniospres globose, dull golden, 23-29 x 21-25 um, wall 2.3-3.7 um
thick, finely to compactly echinulate, 2-3 equatorial germ pores. Telia
epiphyllous, dark brown, compact. Teleutospores, brown, broadly
ellipsoid, 23-32 x 18-27 um, wall 2.4-3.8 um thick, 4.5-7.5 um at apex,
smooth, pedicels yellowish, 95 um.

P. repens is the main host for this rust species, known from Africa
and tropical and eastern Asia. It was reported from Egypt by Reichert
(1921). Later the same specimen has been mentioned by Melchers
(1931) and El-Helaly et al. (1966).

301

Uromyces rumicis (Schum.) Wint. Hedwigia 19:37 (1880).

On Rumex dentatus L. (Polygonaceae), II+IIl.

Localities: Univ. Mansoura farm, Gamasa, Kafr Saad El-Balad.
Uredinia amphigenous, scattered, minute, round, cinnamon.
Urediniospores globoid to ellipsoid, pale brown, 21-29 x 19-25 um.;
wall brown, 1.5-2 um thick, echinulate, 3 equatorial germ pores. Telia
brown, round, scattered. Teliospores subgloboid to ovoid, brown, 25-
34 x 19-27 um; wall dark brown, 2-2.5 um thick, smooth, with
hemispherical papilla; pedicels thin, hyaline, deciduous.

U. rumicis was first collected in Egypt by v. Thumen (cf. Reichert
1921), and later mentioned by Melchers (1931) and El-Helaly ef al.
(1966). It is common in Europe and Asia, rare in the Americas and
Australia. In Africa it has been reported from the Mediterranean area to
S. Africa.

Uromyces setariae-italicae Yosh., Bot. Mag. Tokyo 20:247 (1906).
On Setaria verticillata (L.) Beauv. and S. glauca (L.) Pal. (Poaceae),
I+.

Locality: Univ. Mansoura farm.

Uredinia amphigenous, cinnamon-brown, elongated, scattered.
Urediniospores ellipsoid, 26.5-35 x 24-27.5 um, wall 1.5-2 um thick,
reddish brown, echinulate, 3 equatorial germ pores. Telia amphigenous,
dark brown. Teliospores globoid to obovoid, 18-24 x 16.5-19.5 um,
wall reddish brown, 1-1.5 um thick, smooth, pedicels colorless, thin-
walled, up to 20 um long.

This rust, widespread on many genera of Poaceae in warmer areas,
has previously been reported on S. viridis (L.) Beauv. in Egypt
(Melchers 1931). The two hosts mentioned above are new to this rust in
Egypt, but they are reported as hosts from other African countries.

Uromyces trifolii-repentis Liro & Liro, Bidr. Kanned. Finl. Nat.
Folk 65: 94 (1908) var. trifolii-repentis.

On Trifolium resupinatum L. (Fabaceae), II+III.
Localities: Kafr El-Batikh, Damietta-Ras El-Bar Rd.

302

Uredinia amphigenous, cinnamon-brown. Urediniospores globoid, 19-
24 x 22-26 um; wall light cinnamon-brown, 1.5-2 um thick, echinulate,
2-5 scattered germ pores. Telia hypophyllous, chestnut-brown.
Teliospores globoid, 12.5-21 x 21-30 um; wall cinnamon-brown, 1.5-2
um thick with hyaline papilla, smooth; pedicels colorless, short, fragile.

U. trifolii-repentis is a widespread rust species, reported from Egypt
by El-Helaly et al. (1966) and Elarosi et al. (1974) on T. alexandrinum
L. and T. resupinatum, respectively. In Africa it has been reported on
Trifolium spp. from Morocco and Algeria, and also from S. Africa.

BIBLIOGRAPHY

BISBY, G.R. & P.O.WIEHE. 1953. The rusts of Nyasaland. Mycol.
Papers, No. 54.

CASTELLANT, E. & R. CIFERRI. 1937. Prodromus Mycoflora
Africa orientalis italicae. Bibl. Agr. Colon. Firenze.

CASTELLANIT, E. & R. CIFERRI. 1950. Mycoflora Erythraea,
Somalia et Aethiopica. Supplementum I. Suppl. Atti Ist. Bot.

Univ. Pavia, Ser. 5, vol. H.

CUMMINS, G.B. 1971. The rust fungi of cereals, grasses and bamboos.
Berlin-Heidelberg-New York.

CUMMINS, G.B. 1977. Nomenclatural changes and new species in the
Uredinales. Mycotaxon 5: 398-408.

ELAROSI, H., S.-H. MICHAIL & M.A. EL-MELEIGI. 1974.
Occurrence of rusts of Persian clover and Malva spp. in Egypt.
Alex. J. Agric. Res. 22: 437-441.

EL-HELALY, A.F., 1.A. IBRAHIM, M.W. ASSAWAH, H.M.
ELAROSI, M.K. ABO-EL-DAHAB, S.H. MICHAIL, M.A.
ABD-EL-REHIM, E.H. WASFY & M.A. EL-GOORANI. 1966.
General survey of plant diseases and pathogenic organisms
in U.A.R. (Egypt) until 1965. Alex. J. Agric. Res., Fac.

Agric. Res. Bull. No. 15.

FAHMY, T. 1927. Egypte: Liste complimentaire des maladies des
plantes observees en 1924-25. Moniteur Intern. Proket. d. Plantes
IBY |

303

GJAERUM, H.B. 1977. Additions to the rust flora of Ethiopia.

Boletin de la Sociedad Argentina de Botanica, 18: 8-16.

GJAERUM, H.B. 1985. East African rusts (Uredinales), mainly from
Uganda 4. On families belonging to Apetalae and Polypetalae.
Mycotaxon 24: 237-273.

GJAERUM, H.B. 1987. Rust fungi in the Canary Islands. Vieraea
i ky a9 Beg ale

GJAERUM, H.B. 1990. African host species of Puccinia cyperi-
tegetiformis (Uredinales). Lidia 3: 3-12.

HENNINGS, P. 1891. Fungi africani. Engl. Bot. Jahrb. 14: 339-373.

HENNINGS, P. (1893). Fungi aethiopico-arabici. Bull. Herb. Boissier
1: 26.

JORSTAD, I. 1956. Reliquiae Lagerheimianae. African Uredinales.
Arkiv Bot. 3: 563-598.

JORSTAD, I. 1958. Uredinales of the Canary Islands. Skr. Utg. Set
Norske Vidensk-Akad. Oslo. 1. Mat.-Nat. No. 2.

MAIRE, R. 1905. Notes sur quelques champignons nouveaux ou peu
connus. Bull. soc. Mycol. France 21: 3-12.

MELCHERS, L.E. 1931. A check list of plant diseases and fungi
occurring in Egypt. Trans. Kansas Acad. Sci. 34: 41-106.
PARMELEE, J.A. 1960. The fungi of Ontario. I. Uredinales. Canad.

Dept. Agric., Publ. 1080.

REICHERT, I. 1921. Die Pilzflora Agyptens. Engler ‘s Jahrb. 56: 595-
W275 Vat. I-IV.

ROUMEGUERE, C. 1881. Champignons rapportés en 1880 d’un
excursion botanique en Egypte et en Palestine par M. William
Barley. Rev. Mycologique 3: 23-25.

SNELL, K. 1923. Beitrage zur Kenntnis der pilzparazitaren
Krankheiten von Kulturpflanzen in Aegypten und thre
Bekampfung. Angew. Bot. 3: 121-132. (Ref. R.A.M. 3: 323-324).

STEWART, R.B. & D. YIRGOU. 1967. Index of plant diseases in
Ethiopia. Haile Selassie I Univ., Coll. Agric., Exp. Sta. Bull. 39.

THUMEN, F.V. 1879. Fungi Egyptiaci. Collecti per Dr. Georg
Schweinfurth. Grevillea 8: 49-51.

F
ay
Pieris i ia i
APae 3 mone be yea Ae Tk
’ ‘i
ae ' Ly
“ IF ) ‘
*% i A
ih sti Lice i
‘y ‘ , EL A | cl
} '
i * mo
h * t
i

FORTIS TAT HL SS

ye
Lo ale
12 vay 1

“+:

tke Me 4 he m i
LW adl SUN fT ee a ae BOOK,

ny 1
( , Ly
/ vie j iy * ,
“i GPa Pee
a eet eae Sy een: er "
, poh iy EE. oD e Me ee ¢
fife tale Sh ee Cle ol

} us ‘tends ae is mn Yanan

Wea ATR LO NS aD at Bees ae iy

al i. ; if

‘ : yi te he
i , . ’ ‘ pt na) .
ri } § ‘ 5 va ® Ps a
ME Bae bh Cee ee ee ae | Baers
ne

MYCOTAXON
chess) 2s eR OS Do eT

THE GENUS PYROFOMES IN ARGENTINA AND THE
CULTURAL FEATURES OF P. LATERITIUS
(APHYLLOPHORALES)

Jorge E. Wright, Marcela F. Bolontrade
and Alejandra T. Fazio

Departamento de Ciencias Biologicas, Facultad de Ciencias Exactas y
Naturales, Universidad de Buenos Aires.
1428. Buenos Aires. Argentina

Summary

A study of the two species of the genus Pyrofomes so far
found in Argentina viz., P. latevitius and P. perievis was
undertaken, and the cultural features of P. lateritius are
added. These finds are new records for this country and
broaden the known distribution of the species of the genus in
South America.

Introduction

The genus Pyrofomes was created by Kotlaba & Pouzar (1964)
based on Polvporus demidoffii Lév. (Ryvarden, 1991). It is
characterized by a rather massive applanate to hoof-shaped, perennial
fruit-body, yellowish-orange to rusty-orange, dimitic-trimitic context.
with clamped, hyaline, thin-walled generative hyphae and thick-walled,
strongly rusty yellow, cyanophilous skeletals, occasionally with
binding-like hyaline hyphae; thick-walled, truncate, variably dextrinoid,
cyanophilous basidiospores, and absence of cystidia or other sterile
hymenial elements.

Ryvarden (1972) added another species, P. lateritius (Cke.)
Ryv., based on Fomes lateritius Cooke, described from Rio de Janeiro.

306

Brazil (Saccardo, 1888).

The genus is probably related to Perenniporia and to
Loweporus through its truncate spores, but the former has a white
context, whereas the latter has a dark context and both have dextrinoid
skeletal hyphae.

Pyrofomes demidoffii (Lév.) Kotl. & Pouzar (1964), has been
recorded from Jugoslavia, Russia, Pakistan, U.S., Kenya, Ethiopia,
Tanzania and Uganda. P. lateritius (Cooke) Ryv., originally cited
from Rio de Janeiro, Brazil, was transferred to Pyrofomes by Ryvarden
(1972). P.perlevis (Lloyd) Ryv. (Lloyd, 1912; 1915; Ryvarden 1972,
1989) recorded from Brazil, Colombia, Madagascar and Uganda, is
Close to P. lateritius. P. albomarginatus (Lév.) Ryvarden (1972), has
been cited from Nigeria, Angola, Zaire, Pakistan, India, Sri Lanka,
Thailand, Malaysia, Indonesia, Sarawak, the Philippines, and Australia.
P. aurantiacus David & Rajchenberg, was described from French
Guiana, and P. tricolor (W.A. Murnil) E.J.H. Corner, was recorded
from W. Malaysia, India and Singapore (Corner, 1989).

Pyrofomes demidoffii (Lév.) Kotl & Pouzar has been
described in detail by Bondartsev (1971), as Phellinus demidoffii
(Lév.) Bond. & Sing. and by Gilbertson and Ryvarden (1987). It
appears to be fairly common on living Juniperus in western U.S.,
where it has been known as Fomes juniperinus (von Schrenk) Sacc. &
Syd., causing a white trunk rot. It is also known from East Europe
(Macedonia, Crimea, Caucasus) (Pilét, 1936) and the former Asiatic
USSR (Uzbekistan, Siberia) on living Juniperus excelsa and also on
pines and cypress (Bondartsev, 1971). Pilat (1936) states it is a
subtropical species, seeking drier climate regions. The cultural features
of P. demidoffii have been recorded by Campbell (1938), Nobles
(1958, 1965) and Stalpers (1978).

Pyrofomes aurantiacus was placed in the genus by David &
Rajchenberg (1985) because of the colour of the fruitbody, and
purported to be close to Fomes fulvo-umbrinus Bres. trom Brazil,
which was placed by the same authors in the genus Pyrofomes (David
& Rajchenberg, op. cit.). According to David and Rajchenberg (1985),
the latter species is identical with Perenniporia gomezii Rajchenb. &
Wright, described from N.W. Argentina (Rajchenberg & Wright,
1982), which Ryvarden (1988) accepted in Pyrofomes.

Pyrofomes tricolor, found in the Philippines, Malay Peninsula

307

and India, has a trimitic hyphal system, and is composed of four
varieties that differ in the form of the pileus, the pore surface and the
spore size: P. tricolor vat. tricolor, P. tricolor vat. eumorphus, P.
tricolor vat. shoreicola, and P. tricolor vat. indicus. It causes a white
rot of dipterocarp trees (Corner, 1989) and is related to P. perlevis
(Lloyd) Ryv., which has a tnmitic hyphal system, and grows on
deciduous trees.

Pyrofomes albomarginata (Lév.) Ryv., which is closely related
to P. tricolor, has a paleotropical distribution (Ryvarden and Johansen,
1980). Thus, the specimen of P. Jateritius would be the first record for
Argentina, and since we were able to culture it, we herein describe the
basidiocarps of P. lateritius and P. perlevis and the culture of the
former species.

Materials and Methods

Hand sections of the basidiocarp were made with a razor blade
and mounted in Melzer's reactive, a drop of an aqueous solution of
phloxine mixed with a drop of 5% KOH, and lactophenol-cotton blue.

Replicates of stock cultures of P. lateritius on Difco malt-
extract and Difco Bacto-agar were inoculated in Petri dishes on the
same medium, incubated at 25 °C for six weeks and studied every
week according to the protocol of Nobles (1958).

For oxidase reactions, Petri dishes containing gallic or tannic
acid-Difco malt extract agar were inoculated according to the same
procedure (Nobles, op.cit.).

Other enzymatic tests were carried out with tyrosine, paracresol
and guaiacol (Boidin, 1954).

Colors are according to Munsell (1954).

Taxonomy

Pyrofomes lateritius (Cooke) Ryv., Mycotaxon 31: 45-58, 1988.
= Fomes lateritius Cooke, Grevillea 9:12, 1880.

(Figs. 1-3)

Basidiocarp perennial, sessile, demediate-applanate, 20 x 15 x

308

5 cm; pileus surface covered by moss, radially sulcate towards the
margin, dark reddish brown(3/3 SYR), semibrilliant, turning more
yellowish towards the margin. Context zonate, dark reddish (3/6
2.5YR) to reddish (4/8 2.5YR) or lateritic, below which is a radially
extended granular zone, parallel to the context and permeating the
upper portion of the tubes, intercalating among them as thin whitish
threads, formed by thin-walled, easily staining hyphae with granular
contents, 3-8 zm diam.(Fig. 1 A). Pileus with no xanthochroic reaction

Fig. 1.Pyrofomes lateritius, A. hyphae of granular zone, B. thin-
walled generative hyphae; C. thick-walled generative hyphae; D.
skeletal hyphae; E. basidiospores

309

with 5 %. KOH. Context dark at first with 5 % KOH, but later turning
cherry-red. Tubes ferrugineous, indistinctly stratified; tube length 1-2
mm ;pore surface reddish brown (4/4 5YR), covered by a greyish film:
pores 4-5 per mm, 166-225 pm diam.; dissepiments 64-91 um diam.

Hyphal system dimitic, composed of two types of generative
hyphae: i) hyaline, thin-walled, clamped, easily stained, 2-6 pum
diam.(Fig. 1 B) and it) thick-walled, yellowish, branched, sparsely
clamped, 0.8-4 pm diam. with very thin ends (Fig. 1 D). Basidia not
observed. Basidiospores numerous, with the aspect of chlamydospores,
hyaline to yellowish brown, very thick walled, truncate, variously
shaped but mostly ellipsoid-truncate resembling corn grains, 5.2-6.7 x
4-54 wm, IKI-, slightly cyanophilous (Fig. 1 E).

Culture Description

Macroscopic characters: growth slow, covering 1 cm in one
week and 6.5 cm at the end of six weeks, mycelial mat woolly-felty,
margin silky, evident only at the 6" week; pale yellow (8/4 SY) to
yellow (8/6 SY), with yellow patches (8/6 5Y) during the first three
weeks, later exhibiting a white (8/1 SY) margin which turns pale
yellow (8/3 SY) to yellow (7/6 10 YR) towards the inoculum, and pale

Fig, 2. Culture of
Pyrofomes lateritius.
Aspect of mycelial mat.

310

yellow (8/3 SY) to white (8/1 SY) on the inoculum; reverse unchanged
(Fig. 2). Odorless.

Microscopic characters: Advancing zone: with numerous
clamped generative hyphae, 2.6-4.2 ym diam. (Fig. 3 A), after the a3
week thick- walled generative hyphae appear, 2.1-5.7 ym diam., and
skeletals, 2.1-5.7 pm diam.,stained yellow after the 4" week (Fig. 3
B). Region of inoculum with numerous thin-walled generative
hyphae 1.1-2.1 pm diam.; thick-walled generatives 2.1-2.6 pm
diam. and skeletals 1.1-3.2 um diam. after the 4" week: fibrous hyphae

Fig, 3. Microscopic cultural features of Pyrofomes lateritius, A. thin-
walled generative hyphae; B. skeletal hyphae; C. chlamydospores; D.
chlamydospores and generative hyphae with drops, E. uneven thick-
walled generatives.

311

2-5 ym diam. Chlamydospores 4.6-6.3 x 5.2-15.6 pm, present in the
advancing zone and in the inoculum (Fig. 3 C). After the 4™ week,
drops appear in the generative hyphae and chlamydospores, with a
positive reaction in Sudan TI, both in the advancing zone and in the
inoculum (Fig. 3 D), and irregularly distributed uneven thick-walled
generatives, 2.1-2.6 pm diam., with right-angled branches (Fig. 3 E).
Oxidase reaction: tannic acid= +++ with mycelial
erowth, colony 2.5 cm diam., halo 4 cm; gallic acid= +++ with
variable growth, in some cases only on the inoculum, halo 4 cm diam.;
p-cresol= —, without growth; tyrosine= —, with growth (colony 1 cm
diam.),; 0.2 % guaiacol= +++, with slight mycelial growth on the
inoculum, halo 1.6 cm diam., 2 % suaiacol= —, without growth.
Species code: 2. 3. 8. (9). 24. 25. 34. 37. 38. 47. 53. 54.

Material studied: Argentina, Misiones, Iguazu National Park ,
in “Palo Rosa” (Aspidosperma polyneuron) forest, on living trunk of
“incienso” (Myrocarpus frondosus) associated with a white rot. leg.
JE. Wright, 8-IX-1992 (BAFC 33072; cult. n° 2153).

Remarks: Our material matches well the original description,
except that the pores are smaller (7-9 per mm in the holotype according
to Ryvarden, 1988). We did not observe basidiospores on the basidia.

The present collection expands the distribution area of this
species further South in South America, since it has only been recorded
from Rio de Janeiro and Amazonas, in Brazil.

Pyrofomes perlevis (Lloyd) Ryv., Norw. J. Bot. 19: 236, 1972.
= Fomes perlevis Lloyd, Myc. Writ. 4 Letter No 39:2, 1912.

(Figs. 4, 5)

Basidiocarp perennial, solitary, demediate to triquetous, 7 x 3.5
x 4 cm. Pileus glabrous brown (2/2 SYR), greyish brown (1/4 SYR) in
the older parts, with no xanthochroic reaction. Margin velutinate,
orange to orange-brown (4/8 SYR), 2-2.5 mm wide. Context
ferrugineous-brown (4/6 2.5YR), 3.5 cm thick, later turning cherry-red
with KOH, interrupted by tubes parallel to the context, intercalary
with a granular zone composed of thin-walled hyphae with granular

3h2

Fig. 4. Pyrofomes perlevis, A. hyphae of granular zone; B. skeletal
hyphae with secondary septa; C. binding hyphae, D. basidia, E.
generative clamped hyphae; F. basidiospores.

contents, 2.2-7.8 um diam. (Fig 4 A). Tubes concolorous with the pore
surface, 3-4 mm long, variously stratified. Pore surface whitish with
orange dissepiments, 113-231 j1m diam.; pores 2-3 per mm, 349-392
yum diam.

Hyphal system trimitic (Fig. 5) with scant generative clamped

313

10 Hm

Fig. 5. Pyrofomes perlevis. Hymenial elements

hyphae, 2.1-2.6 pm diam. (Fig. 4 E); skeletal hyphae numerous,
brown, with secondary septa and thin ends, thick-walled, 2.6-3.1 ym
diam. (Fig. 4 B); hyaline, thick-walled hyphae, with almost solid lumen,
branched, are interpreted as binding hyphae, 0.5-2.1 pm diam. (Fig. 4
C). Basidia clavate, 4-spored, 11.4-26 x 5.7-7.8 pm (Fig. 4 D). Cystidia
absent. Basidiospores ellipsoid-truncate, resembling corn grains, thick-
walled, dark brown, 5.2-7.8 x 3.6-5.7 um, IKI-, not cyanophilous (Fig.
4 F).

Material studied: Argentina, Formosa, Ingeniero Suarez, on
Prosopis sp. (“espinillo” and “algarrobo”); leg. Norma Guardia; 10-X-
1986 (BAFC 33634).

Remarks: Both species are very close if not identical.
However, they may be distinguished by pore size, viz. 166-225 pum, 4-5
pores per mm in Pyrofomes lateritius, and 349-392 jm, 2-3 pores per
mm in P. perlevis, and by the dissepiments, 64-91 pm in P. lateritius
and 113-231 um in P. perlevis.

The fact that our study is based on very scanty material does
not warrant conclusions concerning this problem.

On the other hand, a study of the holotypes of both Komes
fulvo-umbrinus and Perenniporia gomezti has confirmed that these
epithets apply to the same taxon. However, we are reluctant to accept
their inclusion in the genus Pyrofomes and rather believe they properly
belong in Perenniporia as already suggested by Corner (1989).

The genus Pyrofomes was described as having sessile-not
resupinate-perennial fruitbodies (Kotlaba & Pouzar, 1964), and none of
the original species included was truly resupinate. Although both this
feature and the color of the context may not be decisive taxonomic
characters on their own, when correlated with others, they may delimit
fairly natural groups. If one relies, for instance, on context color for
separation of taxa, then in this group (Pyrofomes, Perenniporia and
Loweporus) one would have to accept three distinct genera,
notwithstanding microstructural similarities.

In our case, both Pyrofomes lateritius and Pyrofomes perlevis
agree with the diagnosis of Pyrofomes in what concerns truit-body
shape, whereas Pyrofomes aurantiacus and Pyrofomes fulvo-umbrinus
do not. Granted that this feature alone is in many cases useless, its
correlation with the microfeatures allow a separation of a rather
homogeneus group which includes P. demidoffi, P. perlevis, P.
lateritius and probably P. tricolor and P. albomarginatus.

Consequently we treat the latter species as members of
Pyrofomes, and consider Pyrofomes aurantiacus and Fomes fulvo-
umbrinus (= Perenniporia gomezii) as members of Perenniporia.

Acknoledgements

We wish to thank Dr.L. Ryvarden (Oslo, Norway) for his identification
of our material, and Dr. Mario Rajchenberg (Esquel, Argentina) for critically
reading the manuscript.

We acknowledge the assistance received from the authorities of the
Administracion de Parques Nacionales, Argentina, for permission to collect in
the Iguazii Nat’! Park and guide service. The Argentine Nat’l Research Council
provided funds for this research.

We also thank Dr. Edgardo Alberté for the photograph of the culture.

This is paper n° 68 of the PRHIDEB.

Literature

Boidin, J. 1954. Essai Biotaxonomique sur les Hydnés résupinés et les

315

Corticiés-Etude spéciale du comportement nucléaire et des mycéliums.
387 p. Thése. Univ. de Lyon, Fac. des Sciences.

Bondartsev, A.S. 1971. The Polyporaceae of the European USSR and
Caucasia. 896 p. Israel Program for Sci. Translation. Jerusalem.

Campbell, W.A..1938. The cultural characteristics of the species of Fomes.
Bull. Torrey Bot. Cl. 65: 31-69.

Corner, E.J.H. 1989. Ad Polyporaceas V. Nova Hedwigia, Heft 96, 218 p.

David, A. & Rajchenberg, M. 1985. Pore fungi from French Antilles and
Guiana. Mycotaxon 22(2): 285-325.

Gilbertson, R.L. & Ryvarden, L. 1987 North American Polypores. Vol. 2.
Fungiflora. Oslo.

Kotlaba, F. & Pouzar, Z. 1964. Preliminary results on the staining of spores
and other structures of Homobasidiomycetes in cotton blue and its
importance for taxonomy. Feddes Repertorium 69 (2): 131-142.

Liovd. C.G. 1912. Svnopsis of the genus Fomes. Mvcol. Writ. 4 Lett. No. 39:2.

Lloyd, C.G. 1915. Synopsis Fomes. Mycol. Writ. 4. p.231.

Munsell Color Co. 1954. Determination of soil color. From U.S. Dept.
Agriculture Handbook 18. Soil Survey Manual.

Nobles. M.K. 1958. Cultural Characters as a guide to the taxonomy and
phylogeny of the Polyporaceae. Can. J. Bot. 36: 883-926.

Nobles, M.K. 1965. Identification of culmres of wood-inhabiting
Hymenomycetes. Can. J. Bot. 43: 1097-1139.

Pilat, A. 1936. Polyporaceae m Kavina, C. & Pilat, A. “Atlas des
“hampignons d’Europe”, t.III. 624 p. Praha.

Rajchenberg, M. & Wright, J.E. 1982. Two new South American species of
Perenniporia (Polyporaceae). Mycotaxon 15: 306-310.

Ryvarden, L. 1972. A critical checklist of the Polyporaceae m tropical East
Africa. Norw. J. Bot. 19: 229-238,

Ryvarden, L. 1988. Type studies in the Polyporaceae 19. Species described by

M.C. Cooke. Mycotaxon 31: 45-58.

Ryvarden L. 1989. Type studies in the Polyporaceae 21. Species described by
C.G. Llovd in Cvclomvees. Daedalea. Favolus. Fomes and
‘exagonia.. Mycotaxon 35: 229-236.

Ryvarden. L. 1991. Genera of Polvpores. Nomenclature and taxonomy.
Synopsis Fungorum n° 5.Fungiflora, Oslo. 363p.

Ryvarden, L. & Johansen ,i. 1980. A preliminary polypore flora of East
Africa. 633p.

Saccardo, P.A. 1888. Sylloge Fungorum. 6: 188. Patavii.

Stalpers, J.A. 1978. Identifitation of wood-inhabiting Aphyllophorales in pure
culture. Stud. Mycol. 16: 1-248.

ig ne ae the
LM ,

Toei a
ay

ne
i ae :
pods tit Pa ah
uy A ai f
; iY 4

Sere

. y" tae | ee eed

MYCOTAXON

Volume LX, pp. 317-321 October-December 1996

TAXONOMY OF FOMITOPSIS RUBIDUS COMB. NOV.
ANJALI ROY

Department of Botany, Visva-Bharati University, Santiniketan-731 235,
West Bengal, India

and

A. B. DE

Department of Botany, Burdwan Raj College, Burdwan-713 104,
West Bengal, India

SUMMARY

Morphological and cultural characters of Polyporus rubidus Berk.
have been studied in detail and in the light of these findings
the taxonomy of the fungus has been discussed. The new
combination Fomitopsis rubidus (Berk.) Roy & De is proposed.

INTRODUCTION

Polyporus rubidus Berk. grows widely in India on dead hardwoods causing
brown rot. The authors made several collections of this fungus and
studied them morphologically and in culture including interfertility
tests. The purpose of this paper is to describe, in detail, P. rubidus
and to discuss its taxonomic satus.

MATERIALS AND METHODS

Observations were based on fresh specimens collected by the authors.
Microscopic characters of the basidiocarps were studied from freehand
sections mounted in 10% KOH and stained with 1% Cotton blue. Cultures
were established from context tissues of these basidiocarps and studied
following the methods of Nobles (1948, 1965) and Nobles et al. (1957).
Oxidase reactions were tested by the Bavendamm method as described by
Davidson et al. (1938). Voucher specimens and cultures are maintained in
the Mycological Herbarium of the Visva-Bharati University (VBMH).

DESCRIPTION OF BASIDIOCARP

Fomitopsis rubidus (Berk.) Roy & De, comb. nov.
Basionym : Polyporus rubidus Berk., Lond. J. Bot. 6 : 500, 1847.

Basidiocarp annual, rarely biennial to perennial, sessile or
effused-reflexed, corky, becoming woody on age, dimidiate to
semicircular, applanate, conchate or some ungulate, imbricate, 2.0-6.0 x
1.0-3.5 x 0.2-0.5 cm; upper surface cinnamon buff to pinkish brown, at

319

first minutely tomentose, soon glabrous, zonate; margin thin, slightly
incurved; context light pinkish brown, fibrous, 1.0-2.5 mm _ thick;
hymenial surface light pinkish brown to pale cinnamon, pores small,
circular to subangular, 6-8 per mm, pore tubes up to 3 mm long.

Hyphal system trimitic. Generative hyphae hyaline, thin-walled, clamped,
1.5-2.5 wide (Fig.1). Skeletal hyphae (Fig.2) hyaline or nearly so,
thick-walled to subsolid, straight or flexuous, 2.5-4.5 um wide. Binding
hyphae (Fig.3) moderately branched, hyaline, thick-walled to solid,
2.5-4.5 pm wide. Basidia (Fig.4) clavate, 4-sterigmate, 10-15 x 3-5 um.
Basidiospores hyaline, thin-walled, smooth, cylindric-ellipsoid,
apiculate, 5.0-7.5 x 3.5-4.0‘pm (Fig.5), amyloid, cyanophilous.

SPECIMENS EXAMINED : VBMH 80701 on dead wood of Bambusa arundinacea
Willd.; VBMH 80702 on dead wood of Tectona grandis L.f.; VBMH 80703 on

log of Shorea robusta Gaertn. f.; VBMH 84704 on dead wood of Artocarpus
I VBMH

lakoocha Roxb.; VBMH 84705 on dead wood of Castanea sativa Mill.;
85705 on dead wood of Dipterocarpus indicus Bedd.; VBMH 86706 on dead
wood of Eucalyptus citriodora Hook.; VBMH 87707 on dead branch of
Michelia sp.; VBMH 89/708 on dead wood of Terminalia tomentosa Wright et
Arn.

DISTRIBUTION IN INDIA : Dehra Dun (U.P.), Calcutta (W.B.), North and

South India.

DESCRIPTION OF CULTURE

Growth characteristics : Growth slow, plates not covered in 6 weeks.
Advancing zone even, hyaline, appressed or raised, aerial mycelium
extended to the limit of growth. Mat white, cottony-floccose to
wooly-floccose to floccose-felty or wooly-felty, with thin strands of
mycelium raising along the wall of petri dish near inoculum. Reverse
unchanged.

Oxidase reactions negative on both gallic acid and tannic acid agars,
colony 4 cm in diameter on the former and 3.5 cm on the latter.
Microscopic characteristics : Advancing zone with hyphae hyaline,
thin-walled, clamped, branched, often producing many short branches
towards the distal ends, 1.5-5.0 pm wide (Fig.6). Aerial mycelium (a)
hyphae as in the advancing zone; (b) fibre hyphae (Fig.7) hyaline,
thick-walled to solid, branched and unbranched, 1.2-3.0 pm wide;
(c)chlamydospores numerous, hyaline, thick-walled, smooth, terminal and
intercalary, amyloid, cyanophilous, 7.5-12.0 x 3.5-8.0 pm (Fig.8).
SEXUALITY : The result of pairing of 20 monosporous cultures obtained
from a fresh basidiocarp (VBMH 80703) shows the species to be bipolar
with the following distribution of mating types among the basidiospores.
A, Ree IAL Tis celts o titel BOK Y TNS Wye aT iS
Ay ? Zee ag O's UE Uh? ek 5 LA hous 20s

SPECIES CODE : 1. 3. 8. 34. 36. 38. 47. 54. 59 [following the system of
Nobles (1965) |.
CULTURES EXAMINED : VBMH 80701, 80702, 80703, 86706, and 89708.

Figs. 1-8. Fomitopsis rubidus. 1-5. Microscopic structures from
basidiocarp. I. Generative hyphae. 2.Skeletal hyphae. 3.Binding hyphae.
4.Basidia. 5.Basidiospores. 6-8.Microscopic structures from culture.
6.Thin-walled, clamped hyphae showing frequent branches towards the
apex. 7.Fibre hyphae, branched and unbranched. 8.Chlamydospores.

320

DISCUSSION

From the foregoing description it is evident that this species possesses
trimitic hyphal system. The hyphal system. i.e. monomitic, dimitic, or
trimitic of a species has been considered to be taxonomically important
in circumscribing genera (Bondartzeva 1961, Teixeira 1962, Donk 1964,
Fidalgo and Fidalgo 1967). These authors would not include two species
with different hyphal systems in the same genus. Thus Polyporus rubidus
Berk. possessing a trimitic hyphal system, can not be placed under the

dimitic genus Polyporus Mich. ex. Fr.

Ryvarden (1977) studied the type specimen of Polyporus rubidus Berk. and
concluded that it was "Trametes feei (Fr.) Pat.” Later Ryvarden and
Johansen (1980) listed P. rubidus as a synonym of T. feei. Ryvarden
(1977) and Ryvarden and Johansen (1980) regarded P. rubidus (=T. feei)
as a member of the genus Trametes Fr. because P. rubidus resembles
Trametes suaveolens (L.:Fr.) Fr., the type species of the genus Trametes
Fr., by the following features : (i)pileus surface tomentose when young
but becomes glabrous when old; (ii)pores circular to angular;
(iii)hyphal system trimitic with clamped generative hyphae; and
(iv)hyaline, thin-walled, smooth basidiospores. But our study reveals
that P. rubidus causes brown rot, gives negative result in oxidase test
and possesses bipolar type of sexuality whereas T. suaveolens causes
white rot, gives positive result in oxidase test and possesses
tetrapolar type of sexuality (Nobles 1965). These differences rule out
the possibility of considering P. rubidus and T. suaveolens as
congeneric.

Critical observations on the morphological, cultural and certain
biological characters of Polyporus rubidus Berk. reveal its close
relationship with Fomitopsis pinicola (Swartz: Fr.) Karst., the type
species of the genus Fomitopsis Karst. Both produce effused-reflexed to
sessile, woody, applanate to ungulate basidiocarps with circular pores.
Both have a trimitic hyphal system with hyaline, clamped generative
hyphae; ellipsoid-cylindric, hyaline, smooth, delicate-walled
basidiospores; and cause brown rot of wood. The cultural characters of
the two are similar as evident by comparing the species code of F.
pinicola which is 1.3.8.32.34.36.38.44.46.54.55.59 with that of P.
rubidus given above. Moreover, both the species give negative result in
oxidase test and show bipolar sexuality. The great number of
similarities between the two species indicate a _ phylogenetic
relationship. Therefore, it is concluded that Polyporus rubidus and
Fomitopsis pinicola are congeneric and the transfer to Fomitopsis is
proposed.

ACKNOWLEDGEMENT

The authors are greatly indebted to Dr. J. Ginns, Curator, National
Mycological Herbarium of Canada, for critically reviewing the
manuscript.

REFERENCES

Bondartzeva, M.A. 1961. A critical review of the most recent
classification of the family of the Polyporaceae. Bot. Zh.
(Leningrad), 46: 587-593 (Translation, Canada Department of
Agriculture No.73250, 1961).

321

Davidson, R.W., W.A. Campbell and D.J. Blaisdell. 1938. Differentiation
of wood-decaying fungi by their reactions on gallic acid or
tannic acid medium. J. Agric. Res. 57: 683-695.

Donk, M.A. 1964. A conspectus of the families of Aphyllophorales.
Persoonia, 3: 199-324.

Fidalgo, 0. and M.E.P.K. Fidalgo. 1967. Polyporaceae from Trinidad and
Tobago. II. Mycologia, 59: 833-869.

Nobles, M.K. 1948. Studies in forest pathology VI. Identification of
cultures of wood-rotting fungi. Can. J. Res., C-26: 281-431.

Nobles, M.K. 1965. Identification of cultures of wood-inhabiting
Hymenomycetes. Can. J. Bot. 43: 1097-1139.

Nobles, M.K., R. Macrae and B.P. Tomlin. 1957. Results of interfertility
tests on some species of Hymenomycetes. Can. J. Bot., 35:
377-387.

Ryvarden, L. 1977. Type studies in the Polyporaceae 10. Species
described by J.M. Berkeley, either alone or with other authors,
from 1844 to 1855. Norw. J. Bot. 24: 213-230.

Ryvarden, L. and I. Johansen, 1980. A preliminary polypore flora of East
Africa. Fungiflora, Oslo, Norway.

Teixeira, A.R. 1962. The taxonomy of the Polyporaceae. Biol. Rev., 37:
51-81.

hep
? ity 4
Pe
pers
AY.

oA rg

Wee te

MYCOTAXON

Volume LX, pp: 323-326 October-December 1996

STRIGULA OCCULTA, A NEW SAXICOLOUS LICHEN
FROM NEW ZEALAND

P. M. McCARTHY
Australian Biological Resources Study, Flora Section, GPO Box 636,
Canberra, A.C.T. 2601, Australia

W. M. MALCOLM
Micro-Optics Ltd, Box 320, Nelson, New Zealand

ABSTRACT
Strigula occulta P. M. McCarthy & Malcolm sp. nov. (lichenized
Ascomycotina, Strigulaceae) is described from siliceous rock in New
Zealand. The new lichen is related to S. taylorii (Carroll ex Nyl.) R. C.
Harris and its allies from the Northern Hemisphere. A key to the
saxicolous species of Strigula in Australia and New Zealand is provided.

INTRODUCTION
Although the species of Strigula Fr. are predominantly foliicolous, in recent years a
growing number of corticolous and saxicolous taxa have been recognised mainly in
southern Europe and Australasia (Roux & Bricaud 1993, McCarthy 1995). In this
paper, a new saxicolous species is described from New Zealand. While S. occulta P.
M. McCarthy & Malcolm is very different from other species known from the region,
it is clearly related to the corticolous S. taylorii (Carroll ex Nyl.) R. C. Harris and a
handful of other species in the Northern Hemisphere. A key is provided to the six
saxicolous species of Strigula known from Australia and New Zealand.

METHODS
Hand sections and squash preparations were examined in water. Measurements of
algae, thalline hyphae, paraphyses, ascospores and conidia were made at x1000
magnification; those of the excipulum and asci were made at x400 magnification.
Ascospore dimensions are presented as mean values with extreme values in
parentheses.

Strigula occulta P. M. McCarthy & Malcolm sp. nov. Fig. 1
Thallus epilithicus, viridis, continuus vel rimosus, 30—80 um crassus. Algae
Trentepohlia. Perithecia semiimmersa vel superficialia, nigra, partim vel plusminusve
omnino thallo tecto. Involucrellum nigrum, 0.23—0.35(—0.4) mm diam., 30-130 um

9
i;

B
ture asc

°

10

nid
i

ia and co
Mature and
20 pm

perithec
aC,
,D=

of thallus,
hematic)
0.1 mm, C

Habit
(semi-sc
5 mm, B

°

A,
um

eci
0

ni

sotype)
Vertical section of perith
D, Ascospores. Scale A =

Fig. 1. Strigula occulta (

325

crassum. Excipulum fuscoatrum, 15—25 um crassum. Centrum 0.17—0.24 mm
diametro. Asci 78-92 x 11-14 um. Ascosporae 1-septatae, 22-32 x 4—6.5 um, septo
vulgo fissae. Microconidia 2—3 x 0.7—1 um.

Type: New Zealand, South Island, Nelson, Brook Stream Track, NZMS 260
O27:343878, 41°19.1'S, 173°17.6'E, alt. 130 m, on tree-shaded siliceous rock in
disturbed lowland forest, 22.x.1995, W. M. Malcolm 2623 (CHR 507219-holotype;
CANB-isotype).

Thallus crustose, epilithic, pale to medium green, determinate, continuous to rimose,
matt to slightly glossy, smooth to minutely uneven, 30-80 um thick, ecorticate.
Photobiont Trentepohlia; cells 8-16 x 6-15 ym. Mycobiont cells 2-5 um wide.
Prothallus thin, dark grey to black. Perithecia numerous, semi-immersed to almost
superficial, usually partly to almost entirely covered by a 20—60 um thick thalline
layer. Perithecial apex dull black, usually rounded. Ostiole inconspicuous.
Involucrellum black, 0.23—0.35(—0.4) mm diam., 30-50 um thick near the apex,
70-130 um thick and spreading at the base, often subtending the excipulum.
Excipulum dark brown to blackish, 15—25 pm thick. Centrum globose to depressed-
ovate, 0.17-0.24 mm diam. Paraphyses simple to very sparingly branched, 1—1.5(—2)
um thick. Periphyses absent. Subhymenium c. 20 um thick. Asci fissitunicate,
elongate-clavate, 78-92 x 11—14 um; apex rounded or subacute; ocular chamber
minute, convex to tubercular, visible only in immature asci. Ascospores colourless, 8
per ascus, elongate-fusiform, 1-septate, 22-32 x 4—6.5 um, constricted at the septum,
lacking appendages; apices usually bluntly pointed, occasionally rounded, rarely
apiculate; cells similar or distal cell longer and slightly broader than proximal cell,
usually separating at or after dehiscence; demispores (10.5—)14(-18) x (4-)5(-6.5) um
(100 measured); contents clear or guttulate. Conidiomata numerous, semi-immersed,
black, 60-120 um diam. Microconidia elongate-ellipsoidal to fusiform, 2-3 x 0.7—1
um. Macroconidia not seen.

Notes: Strigula occulta is somewhat similar to the corticolous S. taylorii and its allies
from Europe and North America, all of which have a well-defined thallus, small
perithecia and elongate 1-septate ascospores that frequently break at the septum
(Swinscow 1963, Roux & Bricaud 1993). However, the new species is distinguished
by its saxicolous thallus, broader perithecia that are partly or almost entirely
overgrown by the thallus, dark excipulum, discontinuously longer asci, broader
ascospores and smaller microconidia.

Strigula occulta is known only from its type locality at the north end of the South
Island of New Zealand. It grows on tree-shaded siliceous rocks on a steep east-facing
slope in humid, disturbed, lowland forest which is dominated by Hedycarya arborea,
Coprosma grandifolia and Alectryon excelsus in the overstorey. It is associated with
Pannaria immixta, Porina guentheri, P. leptalea, P. partita, an undescribed Porina
sp. (McCarthy & Malcolm, in prep.), Leptogium sp. and several unidentified mosses
and liverworts. The specific epithet is derived from the Latin occultus (hidden), in
reference to the perithecia which are frequently covered by a layer of thallus.

326

KEY TO THE SAXICOLOUS SPECIES OF STRIGULA IN AUSTRALIA AND

NEW ZEALAND
la Ascospores with transverse septa only 2
1b Ascospores muriform 5
2a Ascospores 1-septate B
2b Ascospores mostly 3—7-septate 4

3a Perithecia 0.15—0.28 mm diam. Asci 45-58 x 6-8 um. Ascospores 6-10 x 2-3.5
um; cells not separating at maturity [NE Australia]
S. minutula P. M. McCarthy
3b Perithecia 0.23-0.4 mm diam. Asci 78-92 x 11-14 um. Ascospores 22-32 x
4—6.5 um; cells often separating at maturity [New Zealand]
S. occulta P. M. McCarthy & Malcolm

4a Ascospores (1—)3(—5)-septate, 15.5—26.5 x 4.5-8.5 um [12-20 x 3-6 um in S.
affinis (Roux & Bricaud 1993)]. Macroconidia 3-septate, 12-21 x 2.5-3.5
um. Thallus calcicolous [New Zealand]

S. aff. affinis (Massal.) R. C. Harris

4b Ascospores 7-septate, 22-33 x 5.5-7.5 um. Macroconidia 5—7-septate,
15-25 x 3-4.5 um. Thallus silicolous [E Australia, New Zealand]
S. stigmatella var. alpestris (Vezda) Coppins

5a Perithecia 0.32—0.58 mm diam. Ascospores submuriform, 23-36 x 7—-11.5 um
[NE Australia, Lord Howe Is., New Zealand]
S. australiensis P. M. McCarthy

5b Perithecia 0.42—0.82 mm diam. Ascospores muriform, 37-63 x 10-19 um [New
Zealand] S. johnsonii P. M. McCarthy

LITERATURE CITED
McCarthy, P. M. (1995). New saxicolous species of Strigula Fr. (lichenised
Ascomycotina, Strigulaceae) from Australia and New Zealand. Muelleria 8,
323-329.
Roux, C. & Bricaud, O. (1993). Studo de la genro Strigula (Lichenes, Strigulaceae) en
S-Francio. Graveco de la makrokonidioj. Bull. Soc. Linn. Provence 44, 117-134.
Swinscow, T. D. V. (1963). Pyrenocarpous lichens: 5. Lichenologist 2, 167-171.

MYCOTAXON

Volume LX, pp. 327-360 October-December 1996

XYLARIA SPECIES FROM PAPUA NEW GUINEA: CULTURAL AND
ANAMORPHIC STUDIES

Katleen Van der Gucht

Botany Laboratory, Department M.S.E., University of Gent, K.L.
Ledeganckstraat 35, 9000 Gent, Belgium

ABSTRACT

Cultural and anamorphic characteristics are described and illustrated for 15 Xylaria
taxa collected in Papua New Guinea. For 6 species the anamorph as found in
nature is described and compared with the anamorph as obtained in culture. Based
on these cultural and anamorphic studies Xylana allantoidea, X. badia, X. cubensis,
X. laevis and X. poitei seem to be closely related and belong to the X. cubensis
group; X. multiplex shows great similarities with X. grammica, X. cf. pallida and X.
papulis; X. luteostromata appears to be allied to X. feejeensis.

INTRODUCTION

Members of Xylaria are frequently encountered in their asexual states in the field
and are commonly isolated as endophytes (Rodrigues et al. 1993; Callan & Rogers
1993). These states can be identified only after comparison with anamorphs and
cultures that originated from identified teleomorphic material. Therefore, well-
established teleomorph-anamorph connections are invaluable as a frame of
reference. In this paper cultural and anamorphic data are presented for 15 Xylaria
taxa in order to complement and supplement our understanding of these fungi.

MATERIALS AND METHODS

Air-dried specimens were used to start cultures, following the collecting procedure
described by Kendrick et al. (1979). The culture technique employed is a slight
modification of the technique used by Callan & Rogers (1990). Most of the cultures
were initiated from multiple ascospores dissected from perithecia that had been
rehydrated with sterile aqua destillata. A few cultures were initiated from small parts
of anamorphs found in nature. For this purpose, a small piece of the conidiome
situated in the conidiogenous region, including conidia, was used. Cultural
characteristics were studied on colonies growing on 9 cm diam plastic Petri dishes

328

containing oatmeal agar (Difco), unless another type of agar is indicated in the
description. The plates were incubated at ca. 20°C under alternating daily periods
of 12 h light and 12 h darkness. Whenever possible, the anamorph derived from
cultures was compared with the anamorph as found and collected in nature. Colour
designations used throughout the taxonomic descriptions are from Kornerup &
Wanscher (1978), and appear in parentheses following the colour names.

Dried cultures are retained in the herbarium (GENT) together with the specimens
from which they are derived. A collection of living cultures was deposited in MUCL
(Mycothéque de l'Université Catholique de Louvain-La-Neuve).

Xylaria allantoidea (Berk.) Fr., Nova Acta Regiae Soc. Sci. Upsal. (ser. 3) 1: 127
(1851). - Pl. 1a-c, Pl. 2a-c.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 3-4 weeks, at first white to light orange-
white (6)(A)(2) with plumose margins, later becoming velvety, often with faint
concentric zones, orange-white to brownish grey (4)(E)(2) towards the centre.
Reverse yellowish brown to dark brown towards the centre (5)(D)(4). Exudate
colourless to light orange. HYPHAE parallel, sparingly branched, septate, hyaline,
smooth, locally becoming dark brown, closely septate, becoming irregularly
branched, and finally breaking into segments. STROMATA developing after 1 month,
especially abundant at the colony margin, usually cylindrical to somewhat
flabelliform, at first light orange-white with a brownish orange (5)(C)(3) base, in the
somewhat broadened uppermost half of the stromata becoming grey (8)(B)(1) due
to conidia production, up to 5 mm high x 1-4 mm diam., with the apex remaining
sterile and proliferating. CONIDIOPHORES in palisades, repeatedly branched, hyaline
to subhyaline, smooth, 2-3 um diam. CONIDIOGENOUS CELLS terminal, cylindrical,
hyaline, smooth, 30-40 x 2.5-3 um, bearing in the upper part small circular refractile
conidial scars. CONIDIA acropleurogenous, obovoid to ellipsoid, with a flattened
circular abscission scar at the base, hyaline, smooth, 4.5-6.5(7) x (2.5)3-3.5 um.
COLONIES grew slowly when started from a piece of the anamorph found in nature:
up to 5.5 cm diam. in 3 weeks, greyish orange (6)(B)(3) and concentrically zoned, in
the centrum covered with white cottony mycelium. STROMATA, CONIDIOPHORES,
CONIDIOGENOUS CELLS and CONIDIA as described from the cultures derived from
ascospores.

ANAMORPH AS FOUND IN NATURE

Anamorph formed on stromata distinct from the teleomorphic stromata.
ANAMORPHIC STROMATA cylindric-fusoid, pale yellow to greyish yellow (4)(A-B)(3), up
to 3.5 cm high x 1-2 mm diam. Conidium-bearing regions covering the entire
surface of the stromata. CONIDIOPHORES, CONIDIOGENOUS CELLS and CONIDIA as
described from conidiomata produced in culture.

For description of corresponding teleomorph see Van der Gucht (1995).

OTHER DESCRIPTIONS AND ILLUSTRATIONS: Rogers 1984: 921, figs. 6, 8, 9, 24-26

329

Plate 1: Xylaria allantoidea - ANAMORPH as formed in culture (92-1046): a.
conidiophores (scale bar = 10 ym), b. conidia (scale bar = 5 um), c. brown hyphae with
short protuberances as formed in aging colonies (scale bar = 10 ym); Xylaria badia -
ANAMORPH as formed in culture (89-518c): d. conidiophores (scale bar = 10 um), eé.
conidia (scale bar = 5 pm).

330

(anamorph in culture); Callan & Rogers 1990: 346, figs. 8-16 (sub Xylaria cf.
allantoidea (Berk.) Fr.) (anamorph in culture); Rodrigues et al. 1993: 120, fig. 2.
(anamorph in culture).

SPECIMENS EXAMINED AND CULTURED: EAST SEPIK PROVINCE: Watam, 3°55'S & 144°33'E, sea
level, rain forest near the coast, on dead wood, 17.5.1992, Vyverman & Vyverman 92-1046 (culture)
(GENT). Taway, 4°2'S & 144°13'E, elev. 40-60 m, lowland hill forest, on dead wood, 14.5.1992, Van
der Gucht & De Meester 92-886 (culture) & 92-900 (culture) (GENT). Between Marienberg and Bien,
3°58'S & 144°18'E, mixed flood-plain forest near the border of River Sepik, on dead wood, 16.5.1992,
Van der Gucht & De Meester 92-983 (culture) (GENT).

MADANG PROVINCE: Finisterre Range, near Naru River ford on road Wau-Lae, 5°27'S & 145°32’E, elev.
350 m, rain forest, on dead wood, 27.10.1989, Van der Gucht 89-1269 (anamorphic state) (culture of
the anamorph) (GENT). Road to Bunapas, 9 km beyond the bifurcation Bogia-Bunapas, 4°11'S &
144°47'E, sea level, rain forest, on dead wood, 30.10.1989, Van der Gucht & De Meester 89-1392
(anamorphic state) (GENT). Awar, 4°6'S & 144°50'E, sea level, coastal rain forest, on dead wood lying
on the ground, 18.5.1992, Van der Gucht & De Meester 92-1054a (culture) (GENT). Cape Barschtsch,
near lake Babel, 5°4'S & 145°48'E, sea level, coconut plantation, on a dead coconut stem lying on the
ground, 24.5.1992, Van der Gucht 92-1216 (culture) (GENT).

Rogers (1985) suggested X. allantoidea as a probable member of the X. cubensis
group which are characterized by conidia formed upon special anamorphic
stromata that are usually produced earlier in the year than the teleomorphic
stromata. My observations are in agreement with this suggestion in that they
indicate that anamorphs and teleomorphs are produced on independent structures
in this taxon. Indeed, | could not find any remnants of conidial structures on the
young, immature teleomorphic stromata, which were found on the same log with
the anamorphs. However, field observations throughout a full season of
development are necessary before one can be certain that the anamorphic
stromata stay anamorphic.

Further evidence for placing X. allantoidea in the X. cubensis group is found in the
cultural characteristics. Cultures of X. allantoidea strongly resemble those of X.
cubensis in several respects. Growth rate, colour, and the production of both
flabellate and cylindrical stromata are shared characters.

Xylaria badia Pat., J. Bot. (Morot) 5: 319 (1891). - PI. 1d-e, Pl. 2g-h.

CULTURAL CHARACTERISTICS
COLONIES reaching edge of a 9 cm diam Petri dish in 3 weeks, at first thin, hyaline,
appressed, the centre becoming grey (28)(C-D)(1) and furrowed, at the margins with

Plate 2: Xylaria allantoidea - ANAMORPH as formed in culture (92-1046): a. culture
started from ascospores (9 cm diam. Petri dish), b-c. conidiomata (x 1.5); Xylaria poitei -
ANAMORPH as formed in culture (92-998): d. culture (9 cm diam. Petri dish), e. cylindrical
stromata bearing conidia (x 4), f. minute subglobose stromata bearing conidia (x 3);
Xylaria badia - ANAMORPH as formed in culture (89-518c): g. culture (9 cm diam. Petri
dish), h. conidiomata (x 6).

331

332

patches of white thicker mycelium. Exudate orange. Reverse golden brown (5)(D)(6)
to yellowish brown (5)(E)(4). HYPHAE sparingly branched, septate, hyaline, smooth,
+ 2 um diam. STROMATA formed after 3 weeks at the margins of the colony,
cylindrical, minute, up to 4 mm high x 1-2 mm diam, at first white, becoming
brownish grey (5)(D)(2). Conidium-bearing regions covering the entire surface of the
stromata except for the apex which remains sterile. CONIDIOPHORES in palisades,
dichotomously branched at the base, brown, smooth, up to 25 um long x 2-2.5 um
diam. CONIDIOGENOUS CELLS terminal, cylindrical, subhyaline to light brown, smooth,
12-16 x 2.5-3 um, bearing denticulate conidial scars apically and laterally. CONIDIA
acropleurogenous, obovoid, with a concave circular abscission scar at the base,
subhyaline, smooth, 3.5-4.5 x 1.8-2 um.

For description of corresponding teleomorph see Van der Gucht (1995).

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Hansa Bay, Laing Island, 4°10'S &
144°52'E, sea level, coralligeneous island 700 x 100 m, on dead wood of bamboo, 3.10.1989, Van der
Gucht K. 89-518c (culture) (GENT). Finisterre Range, 40 km beyond the bridge over the Gogol River
on road Madang-Lae, 5°28'S & 145°29'E, elev. 500 m, rain forest, on dead wood of bamboo,
12.11.1989, Van der Gucht K. 89-1920 (culture) (GENT).

This fungus resembles X. cubensis in general appearance. Based on its
anamorphic and teleomorphic characteristics it probably belongs to the X.
cubensis group (Van der Gucht, in press).

X. badia seems to be restricted to bamboo in Papua New Guinea.

Xylaria cubensis (Mont.) Fr., Nova Acta Regiae Soc. Sci. Upsal. (ser. 3) 1: 126
(1851). - Pl. 3a-c, Pl. 4a-f.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 2-3 weeks, at first white with an orange
centre (6-7)(A-B)(3-6), appressed, with faintly zonate white margins, often furrowed,
later darkening from centre outwards, becoming brownish grey to black (6)(E)(2-3)
and covered by lanose or cottony mycelium (white to tan hyphae). Reverse light
orange to greyish orange (5-6)(A-C)(4). Exudate yelowish orange. HYPHAE parallel,
sparingly branched, septate, hyaline, smooth, with age becoming brown, densely
septate and easily breaking apart. Clusters of dark brown, swollen thick walled
cells, remaining in chains are formed in aging colonies. CONIDIOGENOUS
STRUCTURES formed either on small tufts in the centre of the colony, or on
cylindrical to flabelliform stromata. TUFTS formed after 20 days, subglobose,
brownish grey to grey (6)(D-E)(1-2) due to conidia production over the whole surface,
1-2 mm diam. STROMATA developing after 3 weeks, throughout the whole colony (at
surface and periphery of colony), cylindrical or with a flabelliform conidiogenous
upper part, at first white to orange-white with a black base, becoming pink (6)(A)(2)
or grey (5)(B)(6) in areas of conidial production, up to 25 mm high x 1-3 mm diam.
Conidium-bearing regions usually starting in the somewhat broadened uppermost
half of the stromata later covering the entire surface of the stromata. Occasionally

Plate 3: Xylaria cubensis - ANAMORPH as formed in culture (92-1236): a.
conidiophores (scale bar = 10 um), b. conidia (scale bar = 5 ym), c. clusters of swollen
thick walled, dark brown cells, remaining in chains (scale bar = 10 ym); Xylaria poitei -
ANAMORPH as formed in culture (92-998): d. conidiophores (scale bar = 10 ym), e.
conidia (scale bar = 5 um).

334

the stromata remained sterile. No apparent morphological difference between
conidiogenous structures from either location (tufts, stromata). CONIDIOPHORES in
dense palisades, repeatedly branched, hyaline to subhyaline, smooth, 2.5-3 um
diam. Palisades sloughing off in dusty flakes that accumulate in small heaps
underneath the stromata. CONIDIOGENOUS CELLS terminal, cylindrical, hyaline to
subhyaline, smooth, 9.5-15 x 2-3 um, bearing circular refractile conidial scars
apically and laterally. CONIDIA acropleurogenous, obovoid to ellipsoid, with a
flattened circular abscission scar at the base, hyaline, smooth, (3)3.5-5(6) x (1)1.5-
2(3) (avg. + SD: 4.41 + 0.58 x 1.65 + 0.18 um).

COLONIES started from a piece of the anamorph found in nature showed a more
luxuriant growth of the anamorphic stromata. The other characteristics were
identical to the cultures started from ascospores.

ANAMORPH AS FOUND IN NATURE

Anamorph formed on stromata distinct from the teleomorphic stromata.
ANAMORPHIC STROMATA erect, with black, villose bases and flabellate, foliate or
more or less cerebriform, orange-white (5)(A)(2) or greyish (5)(B-C)(1) fertile portions,
3-7 mm total height x 2-8 mm diam. CONIODIOPHORES, CONIDIOGENOUS CELLS and
CONIDIA as described from anamorphic stromata produced in culture.

For description of corresponding teleomorph see Van der Gucht (1995).

OTHER DESCRIPTIONS AND ILLUSTRATIONS: Martin 1970: 95, pl. V:14 (cultural
characteristics); Rogers 1984: 912-921, figs. 1-3, 10-15, 16-20, 27, 28 (anamorph
in culture and nature); Rogers & Samuels 1986: 617-619, fig. 1, 6c (anamorph in
culture); Callan & Rogers 1990: 362, figs. 59 & 60 (sub Xylaria sp. aff. cubensis
(Mont.) Fr.) (anamorph in culture); Rodrigues et al. 1993: 124, fig. 6. (anamorph in
culture).

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Hansa Bay, Laing Island, 4°10'S &
144°52'E, sea level, coralligeneous island 700 x 100 m, S.W. side of the island, 9.5.1992, Van der
Gucht K. 92-817(*) (culture) (GENT). Finisterre range, near Naru River ford on road Madang-Lae,
5°24'S & 145°38'E, elev. 200 m, rain forest, on dead wood, 22.10.1989, Van der Gucht K. 89-968
(anamorphic state) (GENT). North of Alexishafen, between Danip:& Midipur, 5°5'S & 155°46'E, sea
level, secondary disturbed rain forest in association with coconut plantation, on dead decorticated
wood, 2.5.1992, Van der Gucht K. & De Meester L. 92-504 (culture) (GENT). Amron village, 5°7'S &
145°45'E, elev. 40 m, secondary rain forest with lots of palms, on dead decorticated wood, 3.5.1992,
Van der Gucht K. & De Meester L. 92-521(*) (culture), 92-534 (anamorphic state) (culture) (culture from
anamorph), 92-535 (culture) & 92-542 (culture) (GENT). Balek Wildlife Sanctuary, 5°19'S & 145°43’E,
elev. 0-50 m, lowland rain forest, on dead wood, 4.5.1992, Van der Gucht K. & De Meester L. 92-598
(culture), 92-596(*) (culture) (GENT). Awar, 4°6'S & 144°50'E, sea level, lowland rain forest near the
sea with lots of palms, on dead wood, 18.5.1992, Van der Gucht K. & De Meester L. 92-1050(*)
(culture), 92-1052 (culture), 92-1058 (culture), 92-1079 (culture), 92-1086 (culture).

Plate 4: Xylaria cubensis - ANAMORPH as formed in culture (92-1236): a. young culture
started from ascospores (9 cm diam. Petri dish), b. culture started from a piece of the
anamorph, c. subglobose tufts bearing conidia (x 6), d. sterile stroma (x 1.5), e-f.
conidiomata (x 1.5); Xylaria laevis - CULTURAL CHARACTERISTICS (92-1473): g. young
culture (9 cm diam. Petri dish), h. sterile stromata (x 1.5).

335

336

Eo loco, 20.5.1992, Van der Gucht K. & De Meester L. 92-1126(*) (culture), 92-1128 (culture), 92-
1145(*) (culture), 92-1168 (culture), 92-1171(*) (culture). Ramu Region, Brahman, 5°46'S & 145°22'E,
elev. 150 m, disturbed forest near Brahman mission, on dead wood, 25.5.1992, Van der Gucht K. &
De Meester L. 92-1236 (culture) (GENT). Ramu Region, 5°44'S & 145°20'E, elev. 150 m, alluvial flood-
plain forest, on dead wood, 26.5.1992, Van der Gucht K. & De Meester L. 92-1253 (culture) (GENT).
Baiteta, 5°0'S & 145°44'E, elev. 50 m, rain forest with lots of young trees, on dead wood, 2.6.1992, Van
der Gucht K. & De Meester L. 92-1558 (culture) & 92-1562 (culture) (GENT). Eo loco, 2.6.1992,
Vyverman W. & Vyverman R. 92-1592(*) (culture) (GENT).

MOROBE PROVINCE: Lae, 4 km beyond bridge over the Markham River direction Wau, 6°42'S & 146°51'E,
sea level, rain forest, dead wood, 8.11.1989, Van der Gucht K. & De Meester L. 89-1658 (anamorphic
state) (GENT).

The * (see specimens examined) indicates a smaller growth form with the stromata
usually occurring gregariously and characterised by a surface cracked in a
polygonal pattern. This growth form was found in dryer habitats (Van der Gucht
1995). Cultures isolated from both growth forms were identical. These cultures
showed characteristics almost identical with those described in the literature. A
characteristic never mentioned before, however, are the conidial tufts formed in the
centre of several colonies. Such tufts have been observed together with non-
conidial stromata, together with conidium bearing stromata, or without stromata.
The typical Xylocoremium flabelliforme anamorph was found only three times in
nature together with immature or deteriorating teleomorphic stromata. This is an
indication that mature teleomorphic stromata and mature anamorphic stromata
usually do not occur during the same season. The same observation was done by
Rogers (1984). Anamorphic stromata of Xylocoremium flabelliforme are usually
unconnected with the Xylaria cubensis teleomorph. Leessge (personal
communication) indicates, however, that both forms are common simultaneously
in Ecuadorean rain forests.

According to Rogers (personal communication) the presence of pink and of grey
conidial masses, respectively, might indicate that several taxa are, in fact,
represented here, with the cultures having grey conidia produced on stromata
representing X. /Jaevis. Cultures from taxa that | consider to be X. laevis, however,
differ in general appearance (see description of X. /aevis). In any case, many
problems still remain in this group.

Xylaria curta Fr., Nova Acta Regiae Soc. Sci. Upsal. (ser. 3) 1: 126 (1851). - PI.
13h.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 2-3 weeks, at first pure white, velvety with
distinct plumose margins, darkening from centre outwards to olive-brown (4)(F)(5),
later dull black, in the centre becoming overlain by thin white aerial mycelium.
Exudate colourless to light orange. HYPHAE sparingly branched, septate, at first
hyaline, later becoming brown, smooth, + 2 um diam. STROMATA produced within
3 weeks, at first in a concentric zone around the centre, later covering the whole
surface of the colony, abundant, long cylindrical, unbranched, at first white, later
covered from base upwards with villose black hyphae, up to 6 cm long x 0.3-1 mm
diam. The cultures remained sterile.

337

For description of corresponding teleomorph see Van der Gucht (1995).

SPECIMEN EXAMINED AND CULTURED: MADANG PROVINCE: Ramu Region, Road to Bundi, 5°44'S
& 145°20'E, elev. 150 m, alluvial flood-plain forest, on dead wood, 27.5.1992, Van der Gucht K. & De
Meester L. 92-1433a (culture) (GENT).

As has been discussed by Leessge (1987), there have been extensive
misapplications of the name xX. curta. | consider the material collected as true X.
curta, characterized by the presence of the pronouncedly white to yellow
ectostromal squamules (Van der Gucht 1995).

The cultures established by me remained sterile. In general appearance, they
correspond to the description given by Callan & Rogers (1990) sub Xylaria cf. curta
(tropical collection).

The descriptions of cultural characteristics given by Rogers (1983) and Callan &
Rogers (1990) sub Xylana curta are best considered to be cultures of X. comiformis
(Fr.) Fr. fide Callan & Rogers (1993). They differ from the above-mentioned
cultures in that they lack distinctive plumose margins and ‘in that conidiogenous
structures are produced in culture.

Xylaria feejeensis (Berk.) Fr., Nova Acta Regiae Soc. Sci. Upsal. (ser. 3) 1: 128
(1851). - Pl. 5a-c, Pl. 13f-g.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 3 weeks, at first white, very thin with fine
texture and a distinct margin, with centre becoming faint yellow (4)(A)(1-2), later
darkening to reddish grey (7)(B)(2), and becoming furrowed with radiating
depressions, floccose. Reverse pale yellow (4-5)(A)(3-4). Exudate colourless or
yellow to orange. HYPHAE sparingly branched, septate, highly coiled or straight, at
first hyaline, later becoming brown, + 2 um diam. STROMATA formed after 3 weeks
at the margin of the colonies, cylindrical, branched, at first white, later covered from
base upward with villose black hyphae, 5-15 mm high x 2-3 mm diam. Conidium-
bearing regions developing on upper part of stromata, which turns pale tan (4)(C)(3)
due to conidial production. CONIDIOPHORES in dense palisades, branched
dichotomously, hyaline, smooth, 2-3 um diam. CONIDIOGENOUS CELLS terminal,
cylindrical, hyaline, smooth, 8-12 x 2-3 tum, bearing circular refractile conidial scars
apically and laterally. CONIDIA acropleurogenous, obovoid to ellipsoid with a
flattened circular abscission scar at the base, hyaline, smooth, (7)7.5-9(9.5) x 4-4.5
um. Conidia pale tan in mass, often accumulating in conspicuous heaps beneath
stromata.

For description of corresponding teleomorph see Van der Gucht (1995).
OTHER DESCRIPTIONS AND ILLUSTRATIONS: Martin 1970: 120, pl. VI:7, VIl:13

(anamorph in culture); Callan & Rogers 1990: 358, figs. 30-32 (anamorph in
culture).

338

SPECIMEN EXAMINED AND CULTURED: MADANG PROVINCE: Awar, 4°5'S & 144°50'E, sea level,
coastal woodland (with lots of palms and lianes), on dead tree lying on the ground, 20.5.1992, Van der
Gucht K. & De Meester L. 92-1137 (culture) (GENT).

There are some differences in the characteristics of the cultures established by me
compared to data in the literature (Martin 1970; Callan & Rogers 1990). The conidia
of the Papua New Guinean material are larger (7.5-9 x 4-4.5 um) than those
measured by Martin (1970; 6.2-7.5 x 3.1-3.7) and by Callan & Rogers (1990; 5-6
x (2)3 um), and they were produced on cylindrical stromata with black villose bases
instead of on pulvinate stromata (prior to the production of villose basal hyphae) or
on small tufts independent of the stromata as oberved by Callan & Rogers (1990).
However, this latter difference may be only a variation in conidial production. My
own experience with Xy/ara cultures, indeed, indicates that variation may exist
among the kinds of conidiomata produced. One species may produce different
types of conidiomata ranging from small tufts directly on the mycelium to cylindrical
stromata (e.g. X. cubensis).

None of the former studies mentions the striking coiled hyphae of the mycelium.

Xylaria frustulosa (Berk. & M.A.Curtis) Cooke, Grevillea 12: 5 (1883). - Pl. 6a-b.

CULTURAL CHARACTERISTICS

COLONIES on malt extract agar reaching a diameter of 2.5 cm in 4 weeks, at first
white, with a distinct margin, floccose with local aggregations of mycelium, later
becoming greyish brown (6)(D-E)(3). Reverse pale orange (5)(A)(3) to brown (6)(E)(4).
Exudate colourless. HYPHAE sparingly branched, septate, hyaline, smooth, + 2 um
diam. CONIDIOGENOUS STRUCTURES produced on minute, cushion-like tufts, formed
in the centre of the colonies after 3 weeks. Tufts villose and brown (5)(C)(3) when
young, becoming brownish grey (5)(D)(2) due to conidial production, 1-1.5 mm diam.
Conidiogenous regions covering the entire surface of the tufts. CONIDIOPHORES in
loose palisades, dichotomously branched, subhyaline to light brown, smooth, 90-
120 um long x 2-3 um diam. CONIDIOGENOUS CELLS terminal, cylindrical, subhyaline,
smooth, 20-25 x 2-2.5 um, bearing the conidial scars apically and laterally. CONIDIA
acropleurogenous, clavate, with a flattened to concave abscission scar at the base,
hyaline, smooth, 6.5-9.5(11) x 2.5-3.5 um (avg. + SD: 8.74 + 1.39 x 2.85 + 0.04

ym).
For description of corresponding teleomorph see Van der Gucht (1995).

OTHER DESCRIPTION AND ILLUSTRATIONS: Jong & Rogers 1970: 851-855, figs. 1-7
(sub Penzigia frustulosa (Berk. & M.A.Curtis) J.H.Mill.) (anamorph in culture).

SPECIMENS EXAMINED AND CULTURED: EAST SEPIK PROVINCE: Watam, 3°55'S & 144°33'E, sea
level, coastal woodland dominated by Casuarina trees, on dead wood, 17.5.1992, Van der Gucht K.
& De Meester L. 92-1022 (culture) (GENT).

Plate 5: Xylaria feejeensis - ANAMORPH as formed in culture (92-1137): a.
conidiophores (scale bar = 10 um), b. conidia (scale bar = 5 um), c. coiled hyphae of the
mycelium (scale bar = 10 um); Xy/aria grammica - ANAMORPH as formed in culture (92-
1371): d. conidiophores (scale bar = 10 ym), €. Conidia (scale bar = 5 um), f. brown hyphae
with short protuberances as formed in aging colonies (scale bar = 10 um).

340

MADANG PROVINCE: Balek Wildlife Sanctuary, 5°20'S & 145°43’E, elev. 150 m, lowland hill forest, on
dead wood, 11.11.1989, Van der Gucht K. & De Meester L. 89-1815 (culture) (GENT). Awar, 4°6'S &
144°50'E, sea level, coastal woodland and remnant forest, on dead wood, 18.5.1992, Van der Gucht
K. & De Meester L. 92-1069 (culture) (GENT).

The cultural and anamorphic characteristics of the Papua New Guinean material
correspond completely with the above-mentioned description.

Xylaria grammica (Mont.) Fr., Nova Acta Regiae Soc. Sci. Upsal. (ser. 3) 1: 128
(1851). - Pl. 5d-f, Pl. 10d.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 2-3 weeks, at first white, with plumose
margins, velvety, becoming light yellow to orange-white (5)(A)(2), sometimes with a
greenish tint (3)(8)(2), finally darkening from centre outwards (5)(E)(2-3), the centre
covered with white cottony mycelium. Reverse brownish orange, yellowish brown,
golden brown to light brown (5-6)(C-D-E)(5-8). Exudate colourless to orange. HYPHAE
parallel, sparingly branched, septate, becoming dark brown, densely septate, more
irregularly branched, and finally breaking down into segments. STROMATA formed
after 3-4 weeks, in a more or less concentric zone around the centre of the colony,
at first yellowish, later darkening from base upwards, greyish to dark olive-green,
apices remaining white, cylindrical, up to 5 cm total height x 3 mm diam.
CONIDIOGENOUS STRUCTURES formed either on subglobose wart-like structures
developed on the cylindrical stromata or on small dense clusters of conidiophores
directly on the mycelium.

- Conidiogenous structures in dense clusters directly on the mycelium:
CONIDIOPHORES irregularly branched, hyaline, smooth, 2.5-3.5 wm = diam.
CONIDIOGENOUS CELLS terminal, cylindrical or more irregularly shaped, swollen,
hyaline, smooth, 12-32 um x 4-6 um, bearing circular refractile conidial scars
apically and laterally.

- Conidiogenous structures on subglobose wart-like structures:

CONIDIOPHORES in palisades, not branched or branched dichotomously, hyaline,
smooth, 4-5.5 um diam. CONIDIOGENOUS CELLS terminal, cylindrical, hyaline,
smooth, 12-15 x 3.5-5 um, bearing denticulate conidial scars apically.

No apparent morphological difference between the conidia from either location.
CONIDIA acropleurogenous, elongated ellipsoid with a flattened circular abscission
scar at the base, and a broad flattened apex, hyaline, smooth, 12-14(15.5) x 3-
3.5(4) um (avg. + SD: 13.15 + 0.88 x 3.30 + 0.27 um).

For description of corresponding teleomorph see Van der Gucht (1995). .

OTHER DESCRIPTION AND ILLUSTRATIONS: Rogers et al. 1987: 161, figs. 41-43
(anamorph in culture).

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Ramu Region, Paiya, 5°47'S &
145°22'E, elev. 150-200 m, rain forest on hill, on dead wood near stream, 26.5.1992, Van der Gucht
K. & De Meester L. 92-1357 (culture) & 92-1371 (culture) (GENT).

342

The cultural and anamorphic characteristics correspond largely to the ones
described by Rogers et al. (1987), except for the size of the conidia which were
smaller [12-14(15.5) x 3-3.5(4) um versus (16)17-21 x (3)4-5 um], and the
development of long cylindrical stromata bearing fertile subglobose wart-like
structures. This might indicate geographic variations among collections identified
as X. grammica.

Xylaria laevis Lloyd, Xylaria Notes 1, Mycol. Writings 5: 8 (1918). - Pl. 4g-h.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of a 9 cm diam. Petri dish in 10 days, at first white, with
plumose margins, velvety, often with faint concentric zones and radiating ridges,
greyish orange (6)(A-B)(2-3) from centre outwards, becoming overlain with a very
thick layer (often reaching the lid of the Petri dish) of coarse lanose mycelium, the
surface of which finally becomes flocculose and lighter in patches. Reverse pale
orange to orange (5)(A)(3-5). Exudate yellowish orange. HYPHAE parallel, sparingly
branched, septate, hyaline and smooth. Clusters of brown, swollen, thick walled
cells, remaining in chains, are formed in aging colonies. STROMATA formed after 20
days throughout the whole colony, most numerous at the centre, robust, branched,
at first white to orange-white (6)(A)(2), becoming covered from the base upward with
dense brownish villose hyphae. The colonies remained sterile.

For description of corresponding teleomorph see Van der Gucht (1995).

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Amron village, 5°7'S & 145°45'E,
elev. 40 m, lowland hill forest with some garden regrowth, on dead decorticated wood, 3.5.1992, Van
der Gucht K. & De Meester L. 92-519 (culture) (GENT). Hansa Bay, Laing Island, coralligenous island
700 x 100 m, 4°10'S & 144°52’'E, sea level, S.W. side of island, on dead decorticated wood, 9.5.1992,
Van der Gucht K. 92-821 (culture) (GENT). Awar, 4°6'S & 144°50'E, sea level, rain forest located near
the sea, on dead decorticated wood, 20.5.1992, Van der Gucht K. & De Meester L. 92-1122 (culture)
& 92-1153 (culture) (GENT). Ramu region, road to Bundi, 5°44'S & 145°19'E, alluvial flood-plain forest,
on dead wood, 27.5.1992, Vyverman W. & Vyverman R. 92-1436 (culture) (GENT). Balek Wildlife
Sanctuary, 5°19'S & 145°43'E, elev. 50m, lowland rain forest, on dead wood, 29.5.1992, Van der Gucht
K. & De Meester L. 92-1473 (culture) (GENT).

This taxon is closely allied to and has been confused with Xylaria cubensis (Van
der Gucht 1995). It differs in cultural characteristics by colonies covered by a very
thick layer (often reaching the lid of the Petri dish) of coarse lanose mycelium
(colonies from X. cubensis are not covered by a very thick layer of coarse lanose
mycelium) and the colonies remained sterile (in cultures of X. cubensis the
anamorph is produced on cylindrical to flabelliform stromata or on tufts on the
mycelium). See also remarks under X. cubensis.

343

Plate 7: Xylaria Juteostromata - ANAMORPH as formed in culture (92-1300): a.
conidiophores (scale bar = 10 um), b. conidia (scale bar = 5 ym), c. coiled hyphae of the
mycelium (scale bar = 10 um).

344

Xylaria luteostromata Lloyd, Large Pyrenomycetes 2, Mycol. Writings 5: 31 (1919)
[as X. luteostroma]. Pl. 7a-c.

CULTURAL CHARACTERISTICS

COLONIES on oatmeal agar at ca. 20°C, under 12 h light and 12 h darkness,
reaching edge of a 9 cm diam. Petri dish in 2 weeks, at first white, very thin with
fine texture, centre becoming faint orange, later with yellow (3)(A)(4), orange-grey
(6)(B)(2), Olive-grey (3)(D)(3) to brownish grey (4)(C-D)(2) areas, and becoming furrowed
with radiating depressions, the centre finally becoming overlain with white floccose
mycelium. Reverse orange in the centre (5)(B)(5), brown towards the margins
(5)(E)(4). Exudate colourless. HYPHAE sparingly branched, septate, highly coiled or
straight, at first hyaline, later becoming brown, + 2 um diam. STROMATA developing
in the centre of aging colonies, small, rudimentary. Conidium-bearing regions
developing on the rudimentary stromata. CONIDIOPHORES in palisades,
dichotomously branched, brown, smooth, up to 75 um long x 4 um diam.
CONIDIOGENOUS CELLS terminal, cylindrical, subhyaline to light brown, smooth, 12-
20 x 2.5-3 um, bearing circular refractile conidial scars apically and laterally.
CONIDIA acropleurogenous, subglobose, with a flattened circular abscission scar at
the base, hyaline, smooth, 5.5-7 x 5-5.5 um (avg. + SD: 6.40 + 0.80 x 5.08 + 0.39).

For description of corresponding teleomorph see Van der Gucht (1995).

OTHER DESCRIPTIONS AND ILLUSTRATIONS: Martin 1970: 112-113 (cultural
characteristics); Rogers et al. 1987: 163-164, fig. 21 (cultural characteristics).

SPECIMEN EXAMINED AND CULTURED: MADANG PROVINCE: Ramu Region, 5°47'S & 145°23'E, elev.
150 m, alluvial flood-plain forest, on dead wood, 26.5.1992, Van der Gucht K. & De Meester L. 92-1300
(culture) (GENT).

Rogers et al. (1987) mentioned that this species is certainly allied to X. castorea
Berk. and X. feejeensis (Berk.) Fr. The cultural and anamorphic features, indeed,
reinforce the concept of a relationship with X. feejeensis (e.g. general appearance,
texture of the colony surface, presence of coiled hyphae).

Xylaria multiplex (Kunze) Fr., Nova Acta Regiae Soc. Sci. Upsal. (ser. 3) 1: 127
(1851) sensu Dennis, Kew Bull. 11 (1956): 416 (1957). - Pl. 9d-f, Pl. 10h.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 2 weeks, at first white, velvety with distinct
plumose margins, later darkening to greenish grey (28)(E)(2) or dull black from centre
outwards, finally in the centre becoming overlain by thin white aerial mycelium.
Reverse olive-grey (3)(E)(2) in the centre, brownish orange (6)(C)(4) towards the
margins. HYPHAE sparingly branched, septate, hyaline, smooth, + 2 um diam.,
becoming dark brown, densely septate, more irregularly shaped, branched and
finally breaking down into segments. STROMATA produced within 3 weeks,
cylindrical, up to 1 cm long x 1 mm diam., covered from base upwards with villose

345

Plate 8: Xylaria cf. pallida - TELEOMORPH (92-1559): a. aSCi (scale bar = 20 ym), b. apical
apparatus (scale bar = 5 ym), C. ASCOSpores (scale bar = 5 um); ANAMORPH as formed in
culture (92-1559): d. conidiophores (scale bar = 10 im), €. conidia (scale bar = 5 um), f.
brown hyphae with short protuberances as formed in aging colonies (scale bar = 10
lm); ANAMORPH as found in nature (89-1045): g. conidiophores (scale bar = 10 ym), h.
conidia (scale bar = 5 um).

346

black hyphae, remaining white at the top. Stromata remaining sterile.
CONIDIOGENOUS STRUCTURES formed on small white tufts, 3-4 mm diam., developing
within 4 weeks. CONIDIOPHORES in palisades, sparingly branched, hyaline, smooth,
2.5-3 um diam CONIDIOGENOUS CELLS terminal, cylindrical, hyaline, smooth, 12 x 2.5
um, bearing circular refractile conidial scars apically and laterally. CONIDIA
acropleurogenous, elongated ellipsoid witn a small flattened circular abscission
scar at the base, and a broadly rounded to flattened apex, hyaline, smooth, 17.5-
18.5 x 2.2-2.5 um.

For description of corresponding teleomorph see Van der Gucht (1995).

OTHER DESCRIPTIONS AND ILLUSTRATIONS: Martin 1970: 119, pl. IV:6, VI:6 (cultural
characteristics); Rodrigues et al. 1993: 127 (cultural characteristics).

SPECIMEN EXAMINED AND CULTURED: MaADANG PROVINCE: Ramu Region, 5°47'S & 145°23'E, well
drained alluvial forest, on dead wood, 26.5.1992, Van der Gucht K. & De Meester L. 92-1370 (culture)
(GENT).

Based on its cultural and anamorphic characteristics X. multiplex seems closely
related to X. grammica.

Xylaria cf. pallida Berk. & Cooke, J. Linn. Soc., Bot. 15: 395 (1876). - Pl. 8a-h, PI.
10e-g.

TELEOMORPH

STROMATA gregarious, unbranched, cylindrical to clavate with rounded fertile apices,
on well-defined, rather long, slender, smooth stipes (6-20 mm high x 1-1.5 mm
diam.), 2-4 cm total height x 0.3-0.4 cm diam. Externally at first white (immature
stromata), becoming silvery greyish brown (7)(F)(3) to finally dull black with age.
Internally white, becoming hollow with age. Texture hard. Surface smooth, with
older stromata dotted with white to beige granular material. PERITHECIA completely
immersed, subglobose, 0.4-0.6 mm diam.; ostioles inconspicuous to slightly raised,
umbilicate. PARAPHYSES filiform, tapering towards the apex, vaguely septate, + 2 um
diam. near the top. ASC! cylindrical, 8-spored, long stalked, 110-120 x 5.5-6 um
(spore bearing part 45-55 um long); apical apparatus cubic to rectangular, 1.6-1.8
um high x 1.6-1.8 um broad, blueing in Melzer's iodine reagent. ASCOSPORES
uniseriate to obliquely uniseriate, inaequilaterally ellipsoid, with broadly rounded
ends, light to medium brown, smooth, 7-8.5(9) x 3-4(4.5) um; germ silit
inconspicuous, straight, on the ventral side of the spore, almost full spore length.

CULTURAL CHARACTERISTICS

COLONIES reaching 7 cm diam. in 3-4 weeks, at first thin, hyaline, appressed, later
the centre becoming orange-white (7)(A)(2) and furrowed, finally with sectors turning
greyish red (8)(B)(5) to dark grey to almost black (8)(E)(1) with an overall woolly to
floccose aspect; margin not distinct. Exudate colourless to light yellow to orange.
Reverse greyish red (7)(B)(3) in the centre, grey (7)(C)(1) towards the margins.

347

Plate 9: Xylaria papulis - ANAMORPH as formed in culture (92-625): a. conidiophores
(scale bar = 10 ym), b. conidia (scale bar = 5 um), c. brown hyphae with short
protuberances as formed in aging colonies (scale bar = 10 ym); Xylaria multiplex -
ANAMORPH as formed in culture (92- 1370): d. conidiophores (scale bar = 10 ym), e.
conidia (scale bar = 5 um), f. brown hyphae with short protuberances as formed in
aging colonies (scale bar = 10 um).

348

HYPHAE sparingly branched, septate, hyaline, smooth, + 2 mm diam. STROMATA
formed after 3 weeks in a concentric zone near the centre, cylindrical, at first white
to orange-white, turning grey from base upward, up to 1.5 cm high x 1-3 mm diam.
The stromata remained sterile. Conidiogenous structures formed after 1 month on
small, reddish grey (7)(B)(2) tufts. CONIDIOPHORES in loose palisades, di- to
trichotomously branched, hyaline, smooth, up to 90 um long x 2-2.5 um diam.
CONIDIOGENOUS CELLS terminal or intercalary, cylindrical, hyaline, smooth, 12-20 x
2.5-3 um, bearing the conidial scars apically or laterally. CONIDIA acropleurogenous,
elongated ellipsoid, with a flattened circular abscission scar at the base, hyaline,
smooth, 9-12 x 2.5-3 um.

ANAMORPH AS FOUND IN NATURE

Anamorph formed as subglobose or wart-like structures present on immature white
teleomorphic stromata, up to 1-2 x 1-5 mm. Conidium-bearing regions covering the
entire surface of the warts, grey (4)(B)(1). CONIDIOPHORES in palisades, unbranched
or branched near the base, subhyaline, smooth, 2.5-3 Um. CONIDIOGENOUS CELLS
terminal, cylindrical, subhyaline, smooth, 16-25(27) x 3-4 um, bearing circular
refractile conidial scars apically and laterally. CONIDIA acropleurogenous, elongated
ellipsoid, with a flattened abscission scar at the base, hyaline, smooth, 11-13(14.5)
x 3-3.5 um.

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Finisterre Range, near Naru River
ford road Madang-Lae, 5°24'S & 145°38'E, elev. 200 m, rain forest, on dead wood, 11.11.1989, Van
der Gucht K. & De Meester L. 89-1783 (GENT). Baiteta, 5°0'S & 145°44'E, elev. 50 m, rain forest, on
dead decorticated wood, 2.6.1992, Van der Gucht K. & De Meester L. 92-1559 (culture) (GENT). Ramu
region, Paiya, 5°46'S & 145°22'E, alluvial rain forest, Vyverman W. & R. 92-1382 (culture) (GENT).

MOROBE PROVINCE: Lae, direction Gawan, 4 km beyond the bridge over the Busu River, 6°37'S &
147°02’E, elev. 300 m, rain forest, on dead wood, 23.10.1989, Van der Gucht K. 89-1040 & 89-1045
(including the anamorphic state) (GENT).

Since the teleomorph of the material brought in culture was not described before,
| here include a description. The material studied corresponds well to the
description and illustration of Xylaria pallida given by Gonzalez & Rogers (1989:
355, fig. 72), except that the ascospores measure slightly smaller: 7-8.5(9) x 3-
4(4.5) um vs (8.5)9-10(11) x 3.5-4.5 um. Because of this slight difference, and
because | have not been able yet to study the type material, | prefer to call this
fungus X. cf. pallida.

Plate 10: Xylaria papulis - ANAMORPH as formed in culture (92-625): a. culture (9 cm
diam. Petri dish), b. Conidiomata (x 3.5) - ANAMORPH as found in nature (89-992): c.
habitus (anamorph formed on wart-like structures developed on immature
teleomorphic stromata) (x 0.7); Xylaria grammica - ANAMORPH as formed in culture
(92-1371): d. stromata (x 1); Xylaria cf. pallida - TELEOMORPH (92-1559): e. habitus (x 2) -
ANAMORPH as found in nature (89-1045): f. habitus (anamorph formed on wart-like
structures developed on immature teleomorphic stromata) (x 1.5) - ANAMORPH as
formed in culture (92-1559): g. culture (9 cm diam. Petri dish); Xylaria multiplex -
ANAMORPH as formed in culture (92- 1370): h. culture (9 cm diam. Petri dish).

349

350

Based on its anamorphic characteristics here described for the first time, this
fungus seems to be closely related to X. grammica, X. multiplex and X. papulis (see
further).

Xylaria papulis Lloyd, Mycol. Writings 6: 1055 (1921). - Pl. 9a-c, Pl. 10a-c.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 2 weeks; at first white, velvety, with faint
concentric zones, becoming canary-yellow to greenish yellow to dark olive-green
(1-2)(E-F)(7-8), finally darkening from centre outwards, olive-black, the centre covered
with white cottony mycelium. Reverse ochre-yellow (3-4)(A-B)(3-5) to deep wine-red,
greenish at the centre. Exudate greenish yellow. HYPHAE parallel, sparingly
branched, septate, hyaline, smooth, + 2 um diam., becoming dark brown, closely
septate, irregularly branched, and finally breaking into segments. STROMATA formed
after 3 weeks in the centre of the colony, subglobose, moriform to cauliflower-
shaped, up to 0.7 x 0.7 cm, olive-brown (5-6)(B-D)(4), villose when young, turning
olive-grey (5-6-7)(D)(3) due to conidial production. Conidium-bearing regions covering
the entire surface of stromata. CONIDIOPHORES in palisades, sparingly branched,
hyaline, smooth, 2.5-3.5 um diam. CONIDIOGENOUS CELLS terminal, cylindrical,
hyaline, smooth, 16-24 x 3-4 um, bearing circular refractile conidial scars apically
and laterally. CONIDIA acropleurogenous, elongated ellipsoid with a flattened circular
abscission scar at the base, and a broad flattened apex which sometimes appears
to be fringed, hyaline, smooth, 11-13 x (2.5)3-3.5 um.

In one of the cultures (92-625) cylindrical stromata were formed after six weeks, pale
pink when young (with an orange exudate), becoming canary-yellow, green and
finally black from base upwards, villose. No conidiogenous structures observed on
the cylindrical stromata.

ANAMORPH AS FOUND IN NATURE

Anamorph formed on subglobose wart-like structures, grey (4)(C)(1), 2-3 mm diam. ,
developed on immature teleomorphic stromata. Conidium-bearing regions covering
the whole surface of the warts. CONIDIOPHORES in palisades, unbranched or
branched near the base, subhyaline, smooth, 3-4 um diam. CONIDIOGENOUS CELLS
terminal, cylindrical, subhyaline, smooth, 16-20 x 3-4 um, bearing in the upper part
small circular refractile conidial scars. CONIDIA acropleurogenous, elongated
ellipsoid, with a flattened abscission scar at the base and a broadly flattened apex,
hyaline, smooth, (9.5)10-12(13) x (2)2.5-3 um (avg. + SD: 11.12 + 0.91 x 2.56 +
0.32 um).

For description of corresponding teleomorph see Van der Gucht (1995).

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Finisterre Range, near the Naru River
ford on road Madang-Lae, 5°24'S & 145°38'E, elev. 200 m, rain forest, on dead wood, 22.10.1989, Van
der Gucht K. 89-992 (including the anamorphic state) (GENT). Balek Wildlife Sanctuary, 5°19'S &
145°43'E, elev. 50 m, rain forest, on dead wood, 4.5.1992, Van der Gucht K. & De Meester L. 92-625
(culture) (GENT). Ramu Region, 5°46'S & 145°22'E, elev. 150 m, alluvial flood-plain forest, on dead
wood, 26.5.1992, Vyverman W. & Vyverman R. 92-1338 (culture) (GENT).

351

Plate 11: Xylaria schweinitzii - ANAMORPH as formed in culture (92-1343): a.
conidiophores (scale bar = 10 um), b. conidia (scale bar = 5 ym), Cc. coiled hyphae as
found in the mycelium (scale bar = 10 ym); ANAMORPH as found in nature (92-1597): d.
conidiophores (scale bar = 10 ym), €. Conidia (scale bar = 5 um).

352

The production of an olive-green pigment in culture is very characteristic for this
taxon. The same obervation was made by Yu-M. Ju and J.D. Rogers culturing
material from Taiwan (J.D. Rogers & Yu-M. Ju, personal communication). In
general cultural characteristics, as well as in the general conidial shape, the
material shows similarities with the anamorph of X. grammica, X. multiplex and X.
cf. pallida. These taxa represent a conspicuous group of species characterised by
a bright colouration of the colonies as well as of the immature stromata, the
production of conidia on wart-like structures, and the shape of the conidia. Rogers
(1985) who, based upon anamorphic characteristics, proposed a subdivision of the
genus Xylaria into 8 groups divided upon 4 sections did not recognise this group.
He recognized a X. multiplex group with the following members: X. apiculata
Cooke, X. arbuscula Sacc., X. multiplex (Kunze) Fr. and X. schreuderiana van der
Bijl. According to my observations, however, X. multiplex is more closely related to
X. grammica, X. papulis and X. cf. pallida than to X. arbuscula and X. apiculata
(Van der Gucht, in press).

Xylaria poitei (Lév.) Fr., Nova Acta Regiae Soc. Sci. Upsal. (ser. 3) 1: 125 (1851).
P|. 2d-f, Pl. 3d-e.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 3-4 weeks, at first white to orange-white
(5)(A)(2), with plumose margins, woolly, later blackening from centre outwards.
Reverse brown (5)(E)(5). Exudate bright red, orange to yellow. HYPHAE parallel,
sparingly branched, septate, hyaline, smooth, 2-2.5 um diam. CONIDIOGENOUS
STRUCTURES formed after 4-5 weeks either on cylindrical to clavate stromata in the
centre of the colony or on minute subglobose stromata at the colony margin.
Cylindrical to clavate stromata, unbranched or branched, at first orange-white, later
the central part grey (5)(B)(1) due to conidial production, up to 1 cm high x 0.4 cm
diam., with an orange-white apex remaining sterile and proliferating. Minute,
subglobose stromata, often laterally flattened, and joined together with
neighbouring stromata, at first orange-white, later becoming grey due to conidia
production. Conidia-bearing regions covering the entire surface of the minute
stromata. No apparent morphological difference between conidiogenous structures
from either location. CONIDIOPHORES in palisades, repeatedly branched, hyaline,
smooth, 3-4 um diam. CONIDIOGENOUS CELLS terminal, cylindrical, hyaline, smooth,
24-28 x 3-4 um, bearing in the upper part circular refractile conidial scars. CONIDIA
acropleurogenous, obovoid, with a concave or truncate apex (which seems to be
an apical corone by S.E.M., fide Rogers & Callan 1986a), and a flattened circular
abscission scar at the base, hyaline, smooth, 6-8 x 2.5-35 um.

For description of corresponding teleomorph see Van der Gucht (1995).

OTHER DESCRIPTION AND ILLUSTRATIONS: Rogers & Callan 1986a: 287-296, figs. 1-
19 (anamorph in culture and nature).

SPECIMENS EXAMINED AND CULTURED: EAST SEPIK PROVINCE: Watam, 4°58'S & 144°30'E, canoe

353

Plate 12: Xylaria sp., aff. arbuscula - TELEOMORPH (92-1080): a. habitus (x 3), b. detail
of the stromatal surface (scale bar = 0.5 mm), C. aSCi (scale bar = 20 um), d. apical
apparatus (scale bar = 5 ym), €. aSCospores (scale bar = 5 ym); ANAMORPH as found in
nature (92-1080): f. conidiophores (scale bar = 10 ym), g. Conidia (scale bar = 5 um).

354

passage to the Sepik River, mangrove woodland, on dead wood, 17.05.1992, Vyverman W. &
Vyverman R. 92-997 (culture) & 92-998 (culture) (GENT).

The cultural and anamorphic characteristics corresponded closely with those
described by Rogers & Callan (1986a), except that orange tones did not occur in
their cultures.

Xylaria schweinitzii Berk. & M.A.Curtis, J. Acad. Nat. Sci. Philadelphia (ser. 2) 2:
284 (1853). - Pl. 11a-e, Pl. 13a-c.

CULTURAL CHARACTERISTICS

COLONIES reaching a diam of 5 cm in 6 weeks, ‘appressed, at first white, with a
plumose margin, faintly zonate, becoming floccose with a pinkish or reddish cast
(5-6)(A)(2), finally darkening towards grey or black from centre outwards. Reverse
brownish orange to tan (5)(C)(4). Exudate pinkish to colourless. HYPHAE parallel,
sparingly branched, septate, straight or coiled, at first hyaline, later becoming
brown, smooth. STROMATA formed after 6 weeks throughout the colony, cylindrical,
1-5 mm height x 1-1.5 mm diam, at first white to pink, becoming brownish orange,
and finally grey due to conidial production. CONIDIOPHORES in palisades, sparingly
branched, densely septate, subhyaline to brown, smooth, 4-5.5 um diam.
CONIDIOGENOUS CELLS terminal, cylindrical, hyaline to light brown, 12-22 x 4-5 um,
bearing circular refractile conidial scars apically and laterally. CONIDIA
acropleurogenous, ellipsoid, with a flattened abscission scar at the base, hyaline,
smooth, (8.5)9-11 x 3-3.5(4) um. Most of the material brought in culture did not
germinate.

ANAMORPH AS FOUND IN NATURE

Anamorph formed on cylindric-fusoid stromata, orange-grey (5)(B)(2), up to 2.5 cm
length x 1-2 mm diam. Conidium-bearing regions on the upper part of the stromata.
CONIDIOPHORES in palisades, unbranched or branched near the base, subhyaline,
smooth, 3.5-4 um. CONIDIOGENOUS CELLS terminal, cylindrical, subhyaline, smooth,
12-16 x 3-4 um, apically and laterally densely covered with circular refractile
conidial scars. CONIDIA acropleurogenous, ellipsoid, with a flattened circular
abscission scar at the base, hyaline, smooth, 6.5-8(8.5) x 3-3.5 um.

For description of corresponding teleomorph see Van der Gucht (1995).

Plate 13: Xylaria schweinitzii - ANAMORPH as formed in culture (92-1343): a. culture
(9 cm diam. Petri dish), b. young stromata (x 5) - ANAMORPH as found in nature (92-1597):
c. habitus (anamorph formed on cylindric stromata) (x 2); Xylaria sp., aff. arbuscula -
ANAMORPH as formed in culture (92-1073): d. culture (9 cm diam. Petri dish) - ANAMORPH
as found in nature (92-1080): e. habitus (anamorph formed on the sterile apex of an
immature teleomorphic stroma) (x 1); Xylaria feejeensis - ANAMORPH as formed in
culture (92-1137): f. culture (9 cm diam. Petri dish), g. conidiomata (x 2); Xylaria curta -
CULTURAL CHARACTERISTICS (92-1433a): h. Culture (9 cm diam. Petri dish).

355

Nines

piace raeepaiionceo

356

OTHER DESCRIPTIONS AND ILLUSTRATIONS: Rogers & Callan 1986b: 395-397, figs.
11-19 (anamorph in culture); Rogers et al. 1988: 142-143 (cultural characteristics).

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Ramu Region, 5°47'S & 145°23'E,
elev. 150 m, alluvial flood-plain forest, on dead wood, 26.5.1992, Vyverman W. & Vyverman R. 92-
1343 (anamorphic state) (culture) (GENT). Baiteta, 5°0'S & 145°45'E, elev. 50 m, rain forest, on dead
wood, 2.6.1992, Vyverman W. & Vyverman R. 92-1597 (anamorphic state) (GENT).

Cultural and anamorphic characteristics of the material collected in Papua New

Guinea are very similar to those described by Rogers & Callan (1986b), except for

the size of the conidia, the conidia of the specimens studied by me being somewhat
larger: (8.5)9-11 x 3-3.5(4) um vs (7)8-9 x 3-4(4.5) um.

At least two earlier Xy/aria binomials are conspecific with X. schweinitzii and much

detailled work remains to be done in the X. polymorpha group (T. Leessg@e, personal
communication).

Xylaria sp., aff. arbuscula Sacc., Michelia 1: 249 (1878). - Pl. 12a-g, Pl. 13d-e, PI.
14a-c.

TELEOMORPH

STROMATA gregarious, unbranched or branched near the base, cylindrical to
spathulate, with an acute sterile apex (up to 0.5 cm), on a long narrow pubescent
stipe (comprising 1/2 - 2/3 of the total height of the stromata x 1-2(3) mm in diam.),
(2)3-6(7) cm total height x 2-5 mm diam. Externally dark brown (8)(F)(4) to black,
initially covered by a white to grey peeling outer layer. Internally white, solid.
Texture fairly hard. Surface cracked into longitudinally oriented minute scales,
occasionally roughened by protruding perithecia. PERITHECIA partially to completely
immersed, subglobose, 0.5-0.8 mm diam; ostioles conspicuous to inconspicuous,
umbilicate, appearing as small hemispherical black discs. PARAPHYSES filiform,
tapering towards the apex, remotely septate, + 2 um diam. near the top. ASCI
cylindrical, 8-spored, long stalked, 150-180 x 6-7 um (spore bearing part 80-90 um
long); apical apparatus cubic to rectangular, 2.5-3 um high x 2-2.5 um broad,
blueing in Meizer's iodine reagent. ASCOSPORES uniseriate to obliquely uniseriate,
inaequilaterally ellipsoid, with broadly rounded ends, brown to dark brown, smooth,
(11)11.5-13.5(14) x 4-5 um; germ slit spiralling, slightly less than full spore length.

CULTURAL CHARACTERISTICS

COLONIES reaching edge of Petri dish in 4 weeks, at first very thin, bright yellow at
the centre (3)(A)(4) with white plumose margins, becoming thick, velvety to floccose,
often concentrically zonate, orange-white (5)(A)(2) to tan to dark olive-brown (4)(F)(3)
towards the centre. Reverse light yellow (4)(A)(4) to brownish yellow (5)(C)(5). HYPHAE
sparingly branched, septate, hyaline, smooth, + 2 um diam. In aging colonies
clusters of brown, thick walled, subglobose cells are formed. CONIDIOGENOUS
STRUCTURES formed either on small tufts in regular, more or less concentric zones
near the centre of the colony, or directly on the mycelium. Tufts at first orange-
white, later becoming olive-grey (2)(D)(2) due to conidial production. No apparent

357

Plate 14: Xylaria sp., aff. arbuscula - ANAMORPH as formed in culture (92-1073): a.
conidiophores (scale bar = 10 ym), b. conidia (scale bar = 5 ym), c. Clusters of brown, thick
walled, subglobose cells (scale bar = 10 um).

358

morphological difference between conidiogenous structures from either location
(mycelium, tufts). CONIDIOPHORES irregularly branched, subhyaline, smooth, 3-4.5
uum diam. CONIDIOGENOUS CELLS terminal, cylindrical or more irregular in shape,
hyaline, smooth, 10-16 x 3.5-4.5 um, bearing conidial scars in the upper part.
CONIDIA acropleurogenous, obovoid, with a flattened circular abscission scar at the
base, hyaline, smooth, 6.5-8 x 3.5-4.5(5) um.

ANAMORPH AS FOUND IN NATURE
Anamorph formed on the apex (up to 1 cm length x 0.65 mm diam.) of an immature
teleomorphic stroma. Conidium-bearing regions over the whole surface of the apex,
yellowish brown to grey-brown (6)(C)(2). CONIODIOPHORES in palisades, unbranched
or branched near the base, light brown, smooth, 3.5-4 um diam. CONIDIOGENOUS
CELLS terminal, cylindrical, hyaline to light brown, smooth, 10-15 x 3-4 um, densely
covered with denticulate conidial scars. CONIDIA acropleurogenous, obovoid, with
a flattened to concave circular abscission scar at the base, hyaline, smooth, (5.5)6-
7(7.5) x 3.5-4.2 um.

SPECIMENS EXAMINED AND CULTURED: MADANG PROVINCE: Awar, 4°6'S & 144°50'E, sea level,
remnant forest (with many Pandanus sp.) at coast, on dead decorticated wood lying on the ground,
18.5.1992, Van der Gucht K. & De Meester L. 92-1061, 92-1066, 92-1073 (culture), 92-1080 (including
the anamorphic state), 92-1090, 92-1092 & 92-1115 (GENT). Eo loco, 20.5.1992, Van der Gucht K.&
De Meester L. 92-1159 (GENT). Ramu region, Paiya, 5°47'S & 145°23’E, rain forest on hill (open and
dry), on dead decorticated wood, 26.5.1992, Van der Gucht K. & De Meester L. 92-1364 (culture)
(GENT).

A similar fungus has been reported from Venezuela [Rogers et al. 1988: 118, sub
'X. arbuscula (not typical)'] and Mexico (Gonzalez & Rogers 1989: 313-314, figs.
13, 14, sub Xylaria sp., aff. arbuscula). These and my records probably represent
the same taxon, which is, according to the above mentioned authors, close to X.
arbuscula (= X. mellisii (Berk.) Cooke). However, there are some differences in
teleomorph (the smaller size of the ascospores and apical apparatus in the present
taxon) as well as anamorph characteristics [entirely black and white colonies with
no additional colours, black stromata with white apices, and smaller conidia (3)4.5-
5(6) x 2-2.5(3) um produced on the upper surfaces of the stromata or in loose tufts
on the surface of the colony, for X. arbuscula (Martin 1970; Rogers et al. 1988;
Callan & Rogers 1990) versus orange-white to tan colonies, and larger conidia
produced on small tufts or in loose clusters on the surface of the colony for the
species studied here].

X. arbuscula is considered to be a species complex which encompasses several
taxa (e.g. Callan & Rogers 1990). Since this species complex is at present poorly
understood, | consider it premature to erect a new taxon, even though the material
studied represents a taxon clearly different from X. arbuscula s.s.

ACKNOWLEDGMENTS

| sincerely thank Prof. Dr. G. Hennebert for acquainting me with naming and
interpreting anamorphic structures. Special thanks also to Prof. Dr. J.D. Rogers,

359

Dr. Yu-M. Ju and Dr. T. Leessge for sharing their expertise on problematic species
and for reviewing the manuscript. | also thank R. Heynderickx for aid with
photography.

Research was supported by grants n° 2.9006.86 and 2.9001.90 of the Belgian
National Fund for Scientific Research.

LITERATURE CITED

Callan, B.E. & Rogers, J.D. 1990. Teleomorph-anamorph connections and
correlations in some Xylaria species. Mycotaxon 36: 343-369.

Callan, B.E. & Rogers, J.D. 1993. A synoptic key to Xylaria species from
continental United States and Canada based on cultural and anamorphic
features. Mycotaxon 46: 141-154.

Gonzalez, F.S.M. & Rogers, J.D. 1989. A preliminary account of Xylania of Mexico.
Mycotaxon 34: 283-373.

Jong, S.C. & Rogers, J.D. 1970. Penzigia frustulosa in culture. Mycologia 62: 851-
855.

Kendrick, B., Samuels, G.J., Webster, J. & Luttrell, E.S. 1979. Techniques for
establishing connections between anamorph and teleomorpnh. In: Kendrick,
B. (ed.), The whole fungus, the sexual-asexual synthesis. Vol. | & Il.
National Museums of Canada, Ottawa. 793 pp.

Kornerup, A. & Wanscher, J.H. 1978. Methuen Handbook of Colour. 3th edition.
Eyre Methuen, London. 252 pp.

Leess@e, T. 1987. Xylaria corniformis reconsidered. Mycotaxon 30: 81-85.

Martin, P. 1970. Studies in the Xylariaceae: VIII. Xy/aria and its allies. J. S. African
Bot. 36: 73-138.

Rodrigues, K.F., Leuchtmann A., Petrini O. 1993. Endophytic species of Xylaria:
cultural and isozymic studies. Sydowia 45: 116-138.

Rogers, J.D. 1983. Xylaria bulbosa, Xylaria curta and Xylaria longipes in
continental United States. Mycologia 75: 457-467.

Rogers, J.D. 1984. Xylaria cubensis and its anamorph Xylocoremium flabelliforme,
Xylaria allantoidea, and Xylaria poitei in continental United States.
Mycologia 76: 912-923.

Rogers, J.D. 1985. Anamorphs of Xylaria: Taxonomic considerations. Sydowia 38:
255-262.

Rogers, J.D. & Callan, B.E. 1986a. Xylaria poite/: Stromata, cultural description,
and structure of conidia and ascospores. Mycotaxon 26: 287-296.

Rogers, J.D. & Callan, B.E. 1986b. Xylana polymorpha and its allies in continental
United States. Mycologia 78: 391-400.

Rogers, J.D., Callan, B.E., Rossman, A.Y. & Samuels, G.J. 1988. Xylaria
(Sphaeriales, Xylariaceae) from Cerro de la Neblina, Venezuela.
Mycotaxon 31: 103-153.

Rogers, J.D., Callan, B.E. & Samuels, G.J. 1987. The Xylariaceae of the rain
forests of North Sulawesi (Indonesia). Mycotaxon 29: 113-172.

Rogers, J.D. & Samuels, G.J. 1986. Ascomycetes of New Zealand. 8. Xylaria. New
Zealand J. Bot. 24: 615-650.

360

Van der Gucht, K. 1995. Illustrations and descriptions of xylariaceous fungi
collected in Papua New Guinea. Bull. Nat. Plantentuin Belg. 64: 219-403.

Van der Gucht, K. Anamorphs and the classification of Xylaria species. In:
Proceedings of the symposium: "Fungal taxonomy and tropical mycology:
quo vadis ?" - 29th June 1994 - Louvain-la-Neuve (in press).

MYCOTAXON

Volume LX, pp. 361-364 October-December 1996

A NEW SPECIES OF HYGROCYBE FROM INDIA
TK ABRAHAM, KB VRINDA AND CK PRADEEP

‘Tropical Botanic Garden and Research Institute, Palode,
Thiruvananthapuram - 695 562, Kerala, India

ABSTRACT

Hygrocybe parvispora sp. nov., 1s described from Kerala, south
India.

During, continuing and ongoing studies of the agaric flora of
Western Ghats of Kerala, we collected an interesting species of
Hyvgrocybe (Fr.) Kummer, that proved to be undescribed and 1s
considered here as a distinct taxon. The morphological
characteristics were drawn from fresh material. All sections were
prepared in 3% KOH. Colour terminology used is that of
Korncrup & Wanscher (1967). The specimens are deposited at
the herbarium of the Microbiology division, TBGRI, Kerala and
part at Kew.

Hygrocybe parvispora sp. nov. Fig. 1, A-F
Etymology: L. parvispora, small spored

Pileus 4-30 mm _ latus, convexo-depressus, aurantius,
hygrophanus, glaber. Lamellae adnatae cum dente decurrente,
aurantius, lamellulis intermixtac. Stipes 1.5-3 cm x 1-3 mm,
centralis, cylindricus, acqualis, fistulosus, aurantius, glaber.
Sporac (3) 4.5-6 x (2) 3-5.2 ym, subglobosae, hyalinae,
inamyloideac. Basidia 34.5-40.5 x 4.5-9 pm, clavata, 4-

363

sporigera. Acics lamellarum sterilis. Cheilocystidia 14-55.5 x
4.5-12 ym, cylindro-clavata, hyalina. Pleurocystidia nulla.
Trama hymenophoralis regularis, hyalina.

Pileus 4-30 mm diam., convex then plano-convex and slightly
depressed at the centre; surface uniformly ‘deep orange’
(6A8/7A5/7A7/7A8/7C8), drying much paler (6A4/5A2/4A3),
strongly hygrophanous, smooth and glabrous, not viscid, not
translucent striate. Lamellae broadly adnate with a decurrent
tooth, concolorous with the pileal surface, up to 3 mm broad,
moderately close with lamellulae of different lenghts. Stipe 1.5-3
em x I[-3 mm, central cylindric, equal, fistulose, surface
concolorous with the pileus, yellowish towards the base, dry,
smooth, glabrous. Context thin, up to | mm thick ‘brownish
orange’ (6C8), becoming paler, composed of thin-walled, hyaline
hyphae, 3-18 pm diam., with clamp-connexions. Spores (3) 4.5-
6 x (2) 3-5.2 pm, subglobose, thin-walled, hyaline, with a distinct
hilar appendage. Basidia 34.5-40.5 x 4.5-9 pm clavate, bearing
four sterigmata (rarely onc). Lamella-edge sterile, cheilocystidia
abundant, hardly swelling after drying, measuring 14-55.5 x 4.5-
12 yun, variously shaped, more often cylindro-clavate, thin-
walled, hyaline. Pleurocystidia absent. _Hymenophoral trama
regular, hyaline, with parallel hyphae, 3-19.5 wm diam.
Subhymenial layer interwoven. Pileipellis a cutis of semi-erect to
erect hyphae, neither agglutinated nor gelatinized, with the
terminal elements measuring 37-94.5 x 6-15 pm, with
conspicuous clamp-connexions. Caulocystidia abundant on the
upper part of the stipe, scattered and rare elsewhere, 37.5-51.5 x
10.5-18.5 pm cylindro-clavate, thin-walled, hyaline.

Scattered on soil.

?
Mig. 1, A-F T/ygrocybe parvispora. A.habit x 1, 3. basidia;
C.spores, D.cheilocystidia, 1!.cuticular elements, I. caulocystidia
Scale bar = 10 um

364

Specimens examined: [ndia, Kerala: TBGRI campus. 21
October 1993, No.73; 29 October 1993, No.142; 19 November
1993. No.403, 28 June 1994, No. 992; 30 June 1994, No. 1002.
1S July 1994, No.1070, 3 August 1994, No.1279. 6 August
1994, No. 1318, 9 August 1994, No.1335; 10 August 1994, No.
1342. 11 August 1994, No.1348; 2 September 1994, No.1476, 7
September 1994, No.1498; 25 October 1994 No. 1818: 3
November 1994, No.1893;, Agasthyamala,; 2 August 1995,
No.2406, TBGRI campus; 7 August 1995, No.2392, 8 August
1995, No.2402 (holotype), 18 November 1995, No. 2717, 21
November 1995, No.2762.

This small species of //ygrocybe is characterized with inthe
section Coccineae Fayod, by the convex-depressed orange pileus,
lamellae and stipe, adnate to subdecurrent lamellae and lack of
any gelatinization of either the pileal or stipe surfaces. It differs
from all related species in the shape and much smaller size of the
spores, sterile lamella-edge and presence of caulocystidia.

ACKNOWLEDGEMENTS

The authors are grateful to the WGDP of the Department of
Planning, and [Economic Affairs, Government of Kerala for the
financial assistance and to Dr.P.Pushpangadan, Director, TBGRI,
for the encouragement and facilities for carrying out this work.
Help rendered by Dr.D.N.Peglar, Royal Botanic Gardens, Kew
england. in confirming the identity of the species and entically
reviewing the manuscript 1s gratefully acknowledged. ‘Thanks
are also due to Dr.K.J. Joseph, Institute of European Languages,
Thrissur, Kerala for the Latin diagnosis.

REFERENCE

Kornerup, A & Wanscher, JEL 1967. Afethuen Handbook of
Colour, London: Methuen & Co. 243 pp.

MYCOTAXON

nn
Volume LX, pp. 365-372 October-December 1996

NATS TRUFFLE AND TRUFFLE-LIKE FUNGI 5: TUBER LYONII
(= T. TEXENSE), WITH A KEY TO THE SPINY-SPORED TUBER
SPECIES GROUPS

JAMES M. TRAPPE, ARI M. JUMPPONEN AND EFREN CAZARES

Department of Forest Science, Oregon State University,
Corvallis, OR 97331-7501

ABSTRACT

As a result of examination of the type of Tuber lyonii Butters and numerous
collections of Tuber texense Heimsch, the latter is synonymized with the former.
The species is redescribed to encompass its known phenotypic variation, and its
range is extended from earlier reports in Texas, Georgia and Ohio to the presently
known range of northeastern Mexico north to New Mexico, then east along the Gulf
Coast to Florida and north to Minnesota, Québec, New York and Connecticut. The
3-layered wall of the asci of T. /yonii is described, and a key is provided to the three
Tuber spp. that have spores ornamented with spines connected by a low reticulum.

INTRODUCTION

In recent years Mdme. Francisca Marzitelli of Ville d’Anjou, Québec sent
us numerous collections of a Tuber sp. from the vicinity of Montreal that initially
appeared to be an undescribed taxon related to Tuber texense Heimsch
(1958). Examination of all collections of T. texense in our herbarium revealed,
however, that the Québec collections fell within the range of variation of T.
texense and represented a substantial northern and eastern extension of the
range of that species. The range extension in turn prompted examination of
other northern collections assigned by past workers to other taxa in the same
species complex. This included an isotype of T. /yonii Butters (1903), a name

“ synonymized with T. rufum Vitt. by Fischer (1938) and with T. candidum Harkn.
by Gilkey (1939). Tuber lyonii not only differs from these two species in
having reticulate spores but also proved to be conspecific with T. texense.

METHODS

Colors of fresh specimens are in general terms. Hand-cut sections of
both fresh and dried material were mounted in water, 5% KOH, Melzer’s
reagent, or cotton blue for standard light microscopy. Measurements of
structures are from mature specimens. Spores were measured in water
mounts: other reagents may cause spores to either shrink or swell (N. S.
Weber, personal communication). Other structures were measured in KOH
mounts. Herbaria in which examined specimens are deposited are
abbreviated according to Holmgren et al. (1990).

366

TAXONOMY

Tuber lyonii Butters, Bot. Gaz. 35: 431, 1903.
=Tuber texense Heimsch, Mycologia 50: 657, 1958.

Figure 1.

Ascomata hypogeous, subglobose to lobed, with shallow to deep
furrows, 4-30 x 7-55 mm, yellowish brown to orange brown or dark reddish
brown, often with dark brown patches, smooth, the furrows scabrous and
white. Gleba gray in youth, becoming light brown as spores mature, marbled
with narrow, white veins that tend to radiate from the base and emerge through
the peridium in surface furrows. Odor pungent.

Peridium 300-500 um thick. Ectal excipulum (peridial epicutis)
20-40 um thick, of tightly interwoven, light yellowish brown hyphae 4-10 um in
diam at septa, the walls up to 2 um thick, occasional to numerous cells inflated
up to 6-10 (-15) um in diam, often forming a narrow surface zone of
pseudoparenchyma. Ental excipulum (peridial subcutis) of tightly
interwoven, hyaline hyphae 2-5 um in diam, the walls 2-3 um thick.

Gleba of ascus-bearing veins (venae internae) of tightly interwoven,
hyaline hyphae 2-5 um in diam at septa, occasional cells inflated up to 15 um;
sterile veins (venae externae) similar but the hyphae tending toward parallel
alignment, in youth loosely arranged but by maturity compact. Asci
nonamyloid, hyaline, stipitate, the sporiferous part ellipsoid to subglobose, 45-
70 (-100) x 35-70 um, abruptly narrowed to a + cylindric, straight to sinuous
stem (10-) 25-60 x 4-6 (-8) um, the base forked; at maturity the walls have
three layers, seen most clearly in cotton blue mounts: outer wall thin,
inconspicuous, membranous, partially evanescent, loose and baggy around
the ascus base; middle layer 3-5 um thick; inner layer thin, inconspicuous,
membranous, tightly appressed to middle layer. Paraphyses absent.

Spores ellipsoid, yellow to yellowish brown, 1-4 per ascus, in 1-spored
asci 30-37 x 22-24 um excluding ornamentation, in 2-spored asci 25-32 x 18-
23 um, in 3-spored asci 21-31 x 17-21 um, in 4-spored asci 19-24 x 15-19 um;
ornamentation of subhyaline spines (1-) 2-4 x 0.3-1 um mostly connected by
lines on the spore surface to form low alveolae with (3-) 4-6 (-7) sides and
numbering 7-12 along the spore length; spore walls 1-2 (-3) um thick. Spores
light brown in Melzer’s reagent, the reticular ridges hyaline, in cotton blue the
walls deep blue, the spines medium blue.

Etymology: in honor of H. L. Lyon, collector of the type.

Distribution, habitat and season: northeastern Mexico northwest to New
Mexico, east along the Gulf Coast to Florida, north to Minnesota, and east to
Connecticut and southern Québec (Fig. 2); a probable mycorrhizal associate
with Carya illinoensis (Wangenh.) K. Koch, C. ovata (Mill.) K. Koch, Crataegus
sp., Quercus coccinea Muenchh., Q. macrocarpa Michx, Q. polymorpha
Schlecht., Q. rysophylla Weath., Q. turbinella Greene and other Quercus spp.,
and Tilia americana L.; June through March.

Figure 1. Tuber lyonii. Fig. 1A: ascomata, vertical cross section
(left), surface view (right), scale bar = 1 cm. Fig. 1B: 2-spored
ascus showing the loose-fitting, thin outer ascus wall and two
completely reticulate spores, scale bar = 20 um. Fig. 1C: spore
with partial reticulum, scale bar = 7 um.

367

368

Collections examined: ISOTYPE—U. S. A., Minnesota: Hennepin Co.,
vicinity of Minneapolis, H. L. Lyon, 3 Mar. 1903 (holotype FH, isotype OSC

58742). OTHER COLLECTIONS: Canada, Québec: Anjou, St. Gotique, F.
Marzitelli, Trappe 11601, 22 Oct. 1990 (OSC 58747); Deux Montaignes, F.
Marzitelli, Trappe 15106, 7 Nov. 1993 (OSC 58751); Lachute, F. Marzitelli,
Trappe 11325, 30 Oct. 1989 (OSC 58746); Montreal, Ville d'Anjou, F.
Marzitelli, Trappe 11650, 4 Aug. 1990 (OSC 58748); Montreal West, leg. F.
Marzitelli, Trappe 12669, 18 Oct. 1992 (OSC 58750); Park Angrignon (west of
Montreal), F. Marzitelli, Trappe 12453, 25 Oct. 1991 (OSC 58749). México,
Nuevo Leon: Municipio de Garza Garcia, Chipinque, E. Cazares & J. Garcia,
24 Sept. 1983 (ITCV as Tuber rufum var. nitidum, OSC 58767, 58774);
Santiago, Camino Camotera-Potrero, E. Cazares 89, 17 Sept. 1983 (ITCV as
Tuber rufum var. nitidum, OSC); Santiago, Cahén de Puerto Genovevo, E.
Cazares, 8 Oct. 1982, 17 Oct. 1972, 13 Nov. 1982, & J. Garcia, 23 Oct. 1983
(ITCV as Tuber rufum var. nitidum, OSC 58758, 58759, 58760, 58761, 58762);
Santiago, Canhén del Tejocote, E. Cazares, 16 Sept. 1983 (ITCV as Tuber
rufum var. nitidum, OSC 58766); Santiago, Col. Los Pescadores, E. Cazares,
8 Oct. 1983 (ITCV as Tuber rufum var. nitidum, OSC 58763); Santiago, El
Barro, J. Garcia, 27 Nov. 1983 (UNL & ITCV as Tuber rufum var. nitidum, OSC
58771, 58772); Santiago, El Cercado, J. Garcia, 8 Oct. 1983 (ITCV as Tuber
rufum var. nitidum, OSC 58773); Santiago, El Ranchito, J. Garcia, 14 Sept.
1983 (ITCV as Tuber rufum var. nitidum, OSC 58768, 58769, 58770);
Santiago, El Salto, Camino Camotera-Potrero, E. Cazares, 18 Sept. 1983
(ITCV as Tuber rufum var. nitidum, OSC 58764). Tamaulipas: Ciudad
Victoria, El Madrono, J. Garcia, 20 Oct. 1988 (ITCV as Tuber rufum var.
nitidum, OSC 58776); La Palma, Carr. Cd. Victoria-Soto la Marina, J. Garcia,
22 Oct. 1984 (ITCV as Tuber rufum var. nitidum, OSC 58775). United
States, Connecticut: Fairfield Co., near Danbury, L. Robba, 29 Nov. 1952
(OSC 58756). Florida: St. Johns Co., St. Augustine, P. L. Dinkins, Gilkey

1186, 8 July 1964 (FLAS, OSC). Illinois: Big Rock, F. D. Merrill, 23 Aug.

1928 (BPI as Tuber rufum, OSC 58778). lowa: Marshall Co., lowa State Univ.
Rhodes Farm, C. Walker, Trappe 6378, 10 Sept. 1981 (OSC 58743).
Mississippi: Oktibbeha Co., W. G. Welles, Trappe 8779, 4 Dec. 1985 (OSC
58744). New Mexico: Eddy Co., Carlsbad, E. E. Staffeldt, Trappe 4079

(ITCV, OSC). New York: Dutchess Co., L. Robba, 24 Nov. 1953 (OSC
58755); Orleans Co., Medina, L. H. Weld, August (OSC 58754); Westchester

Co., L. Robba, 28 Oct. 1950 and 6 Nov. 1951 (NY as Tuber candidum, OSC
58753, 58757). Ohio: Portage Co., near Brady Lake, S. J. Mazzer, 29 Sept.
1975 (CUP 54704 as T. texense, OSC 58777). Oklahoma: Tulsa Co., Tulsa,
Mrs. C. Connors, Gilkey 1161, 21 Sept. 1962 (OSC). Tennessee: Johnson
Co., Butler, Dale Gregg, Gilkey 823, 2 Feb. 1954 (OSC); Knox Co., near
Knoxville, L. R. Hesler 8262, 24 July 1935 (OSC, TENN). Texas: Comal Co.,
Fischer, M. Forbes, Trappe 11146, 2 June 1989 (OSC); Glasscock Co.,
Garden City, R. Taber, Trappe 5975, September 1980 (OSC 58745); Harris
Co., Houston, L. Guenzel, Trappe 5565, 16 Aug. 1979 (OSC); Travis Co.,
Austin, C. Heimsch 1 (OSC, isotype of T. texense). Wisconsin: Kenosha Co.,
Bristol Woods County Park, A. Parker, Trappe 17304, 27 Aug. 1983 (OSC
58752); Waukesha Co., 2 mi. W. of Mukwonago, J. Steinke, Trappe 17689, 17
Aug. 1993 (OSC).

370

Discussion: Tuber texense was previously reported from Texas (Heimsch
1958), Ohio (Host & Mazzer 1982) and Georgia (Hanlin, et al. 1989). The type
of T. lyonii and the other collections we examined strikingly extend the known
range of this species.

We initially regarded the collections from Québec by Mdme. Marzitelli
as a Closely related but different taxon than T. texense, because the peridial
epicutis was pseudoparenchymatous only in places and then not strongly so.
The type of T. lyoniiis also that way. The type of T. texense and other southern
collections, in contrast, typically have a narrow but well developed
pseudoparenchymatous epicutis. However, examination of many collections
revealed this to be a highly variable character, even on individual specimens,
and hence not a distinguishing trait.

The 3-layered ascus walls of T. lyonii were noted by Trappe (1969, as T.
texense), who observed similar structures in the T. rufum Fr. and T. excavatum
Vitt. complexes of species. He suggested that this character is so distinct from
other Tuber spp. that it justified generic separation as Corda’s proposed genus
Oogaster (Zobel 1854). However, Uecker and Burdsall (1977) observed
similar asci in still other groups of Tuber, and that large and diverse genus has
many other intergrading groups. Consequently, we prefer not to recognize the
genus Oogaster unless stronger evidence, especially molecular and
ultrastructural, becomes available on phylogenetic relationships within the
genus Tuber.

Most collections of T. lyonii have spores with consistently well
developed reticula. Spores of many collections (Trappe 8779 from Mississippi
and 11325 from Québec, all Mexican collections cited) contained many spores
with the reticulum incomplete to occasionally lacking. This variability led
Cazares et al. (1992) to interpret the Mexican material as Tuber rufum var.
nitidum (Vitt.) Fischer. Reexamination of those collections in light of our new
data on Tuber lyonii in the United States and Canada, however, revealed that
in these cases many to most spores do have well developed reticula.

ECOLOGY

Little is known of the ecology of T. /yonii. Butters (1903) wrote of the
type locality: “The locality where the specimens were found is a steep bank
with southern exposure. The soil is a stiff calcareous clay, covered with a
shallow layer of leaf-mold. At the time of collection a week of warm weather
had melted the winter’s snow and thawed the soil to the depth of several
inches, but the clay subsoil was still frozen hard, and the surface soil had been
soft for only a very few days. The fungi were found in the leaf-mold about the
base of a small group of basswood trees (Tilia americana L.)...All the fungi
collected were in a mature condition. There had hardly been time for their
development this spring...It seems probable that the ascomata of the truffle are
formed in the late autumn, and lie over the winter in a practically mature state.”

Tuber lyonii seems to be especially common in places under Carya
illinoensis. \n transmitting his specimens (as T. texense) from New Mexico,
Eugene Staffeldt commented: “They were found under pecan trees believed
to be 25 years old. We examined a number of younger trees in this area and
did not find any specimens on trees younger than 10 years old. During the
day we visited 18 residences (most with 2 to 25 trees) and T. texense was
common to most.” Linda Guenzel commented on her Texas collections from

O71

under oak that “... squirrels love them. They were found just below the ground;
the larger ones had pushed their way up to the surface and were visible,
though still somewhat surrounded by soil.” She also noted that grass
appeared to die where the truffles fruited (perhaps the “brulé” or “truffle burn”
phenomenon commonly observed with Tuber melanosporum Vitt.).

This truffle can be locally abundant. Of his Wisconsin collections, Alan
Parker wrote, "I gathered approximately 110 ascocarps...at one location. A
few of the fruiting bodies were partially exposed on bare soil, which led to the
initial discovery; many others were located by shallow digging in the area.
The collection was made in a disturbed area on the edge of an oak-hickory
woods; the fruiting appeared to be associated with a species of Crataegus."

A KEY TO TUBER SPP. WITH SPINY-RETICULATE SPORES

Three Tuber spp. are now known to have spiny spores with the spines
connected by low lines on the spore surface to form a partial to complete
reticulum: T. lyonii, T. spinoreticulatum Uecker & Burdsall (1977), known only
from the type collection in Maryland, and T. malacodermum Fischer (1923),
known only from Germany and Switzerland. These are all closely related to
the common and widely distributed Tuber rufum Fr. vars. rufum and nitidum ,
taxa with spines but no reticulum on the spores. Many collections of this group
have been misidentified, as we discovered in the course of this study.
Accordingly, a key is presented here. Both varieties of T. rufum have a number
of variants, especially in spore shape (Fischer 1923) and peridial structure.
These need further study to determine whether or not they represent different
taxa, so we do not separate them in the key.

ttopores wilh spines’ but noiretic ules; 0 eae av ce tae onweautedvecauonanecct 2
1. Spores with spines and mostly with a low, partial to complete reticulum......3
2. Ascomata distinctly verrucose.................000008 Tuber rufum var. rufum group

2. Ascomata glabrous to scabrou................... Tuber rufum var. nitidum group

3. Surface of ascomata with warts up to 1 mm broad..... Tuber spinoreticulatum
Go oultace: Of ascomata SIMOO LO. SCADTOUS tii iccfeicseravescsstacperacdaiaccetbedsclevecenassecs 4
4. Peridial epicutis with many cells inflated to 6-10 pm................ Tuber lyonii

4. Peridial epicutis of cells 10-25 (-70) um in diam..... Tuber malacodermum

ACKNOWLEDGMENTS

This series is a project of the North American Truffling Society (NATS).
The research was supported by National Science Foundation Grants DEB
9007186 and 9400545 and the U. S. Forest Service, Pacific Northwest
Research Station.

372

Literature Cited

Butters, F. K. 1903. A Minnesota species of Tuber. Bot. Gaz. 35: 427-431.

Cazares, E., J. Garcia, J. Castillo and J. M. Trappe. 1992.
Hypogeous fungi from northern Mexico. Mycologia 84: 341-359.

Fischer, E. 1923. Zur Systematik der Schweizerischen Triffeln aus den
Gruppen von Tuber excavatum und rufum. Verh. Naturforsch. Gesell. Basel
35: 34-50.

Fischer, E. 1938. Tuberineae. /n A. Engler & H. Harms, Die nattirlichen
Pflanzenfamilien, 2nd. ed., Band 5b. VIII. 42 pp.

Gilkey, H. M. 1939. Tuberales of North America. Ore. State Mon. 1: 1-63.

Hanlin, R. T., M.-L. Wu and T. B. Brenneman. 1989. The occurrence
of Tuber texense in Georgia. Mycotaxon 34: 387-394.

Heimsch, C. 1958. The first recorded truffle from Texas. Mycologia 50:
657-660.

Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index
Herbariorum Part |: The Herbaria of the World. 8th Edition. New York
Botanical Garden, New York. 693 pp.

Host, G. E. and S. J. Mazzer. 1982. An Ohio record for Tuber texense.
Ohio J. Sci. 82: 142.

Trappe, J. M. 1969. Laminated ascus walls in the Tuberales. Abstr. 11th
Internat. Bot. Congress, Seattle. p. 221.

Uecker, F. A. and H. H. Burdsall, Jr. 1977. Tuber spinoreticulatum, a
new truffle from Maryland. Mycologia 69: 626-630.

Zobel, J. B. 1854. Augusti Car. Jos. Corda, lconum Fungorum Hucusque
Cognitorum. Tomus VI (Ultimus). J. Cramer Reprint, 1963. 91 pp.

MYCOTAXON

Volume LX, pp. 373-386 October-December 1996

HYPHOMYCETES ON ALCHORNEA TRIPLINERVIA (SPRENG.) MUELL.
ARG. LEAF LITTER FROM THE ECOLOGICAL RESERVE JUREIA-
ITATINS, STATE OF SAO PAULO, BRAZIL.

Rosely Ana Piccolo Grandi
Instituto de Botanica, CP 4005, 01061-970, Sao Paulo, SP, Brazil.

Derlene Silva Attili
Fundag4o Tropical de Pesquisas e Tecnologia “André Tosello”, CP 1889, 13087-010,
Campinas, SP, Brazil.

ABSTRACT

The following sixteen Hyphomycetes isolated from Alchornea triplinervia
leaf litter, sampled at the Ecological Reserve Juréia-Itatins, are described: Chalara
aurea, Chloridium virescens var. virescens, Cladosporium cladosporioides,
Cryptophiale kakombensis, C. udagawae, Cylindrocladium scoparium, Dictyochaeta
simplex, Idriella falcata, Kionochaeta ramifera, Lauriomyces heliocephala,
Mycoleptodiscus brasiliensis, Phaeoisaria clematidis, P. infrafertilis, Subulispora
procurvata, S. rectilineata and Thozetella cristata. This work aims to contribute to the
knowledge on the fungal diversity of an important Brazilian Rain Forest Reserve,
where studies of the mycota are rare.

INTRODUCTION

Hyphomycetes play an important part in the decomposition process in
ecosystems. The leaf litter is one of the most diverse habitats, exhibiting an intense
microbial colonization, which can develop inside the leaf or on its surface (Cooke &
Rayner, 1984). In Brazil there are very few works on saprobic Hyphomycetes in the
local literature. The Ecological Reserve Juréia-Itatins (ERJI), on the south coast of the
State of Sao Paulo, is considered one of the most representative areas of the Brazilian
tropical ecosystems (Cortesdo, 1989). Only three studies on soil fungi from this area
have appeared (Attili, 1994; Chatanon et al., 1994; Garlipp, 1995). In this paper,
Hyphomycetes isolated from decomposing leaves of Alchornea triplinervia are
identified and characterized, also contributing to the knowledge on the geographical
distribution of these taxa.

374

Fig. 1. Ecological Reserve Juréia-Itatins (E.R.J.1), SP.

375

The Ecological Reserve Juréia-Itatins

The study area is located between 24°18’47”-24°36’10” S and 47°00’03” -
47°30’07” W (Fig. 1). Main climatic characteristics are: mean air temperatures
between 35-15°C, high relative air humidity (80-100%), rarely under 50%; high rain
fall (annual average above 3,400mm), and a less-rainy season during autumn-winter
(Attili et al., 1995). Under these conditions there are a great variety of ecosystems
such as dunes, mangrove, “restingas”, lowland forests, luxuriant rain forest, montane
vegetation and elevations (Cortes4o, 1989). The collecting site is known as “Banhado
Grande’, in the middle of the Reserve.

MATERIALS AND METHODS

Decomposing leaves of Alchornea triplinervia, a common member of the
Euphorbiaceae in the ERJI, were used as substrate for the isolation of the
Hyphomycetes. Samplings were carried out in January, April, July and October 1991
and May 1993, resulting in 25 leaf samples. A washing technique with 15 serial
changes of sterile distilled water (Harley & Waid, 1955) was applied to the leaf litter.
Leaf fragments were left in moist chambers at room temperature until the development
of fungal structures. Identifications were made based on slide mountings prepared with
lactophenol cotton-blue or PVL resin mounting media (Trappe & Schenck, 1982).
Preserved specimens were deposited in the “Herbario Cientifico Maria Eneyda P.
Kauffmann Fidalgo” (SP), at the “Instituto de Botanica”, State of Sao Paulo, Brazil.

RESULTS

The following taxa were isolated during the collecting period:
Chalara aurea (Corda) Hughes in Can.J.Bot., 36: 747, 1958 (Fig. 2).
Conidiophores erect, single or in groups, cylindrical, brown, septate, smooth, 20.0-
67.0 x 5.0-6.5 um. Conidiogenous cells phialidic, integrated, erect, brown, smooth,
cylindrical or lageniform, with gradual transition venter/collarette, 77.5-102.5 x 5.5-
7.0 um. Venter with superior limite slightly pronounced; tubular collarette. Conidia
enteroblastic, catenate, hyaline, cylindrical with truncate ends, 1-septate, smooth-
walled, 15.0-18.5 x 2.0-3.0 um.
Material examined: SP250853.
Literature consulted: Nag Raj & Kendrick, 1975; Holubov4-Jechové, 1984.

Chloridium virescens (Pers.) W. Gams & Hol.-Jech. var. virescens in Stud. Mycol., 13:
17, 1976 (Fig. 3-4).

Conidiophores erect, single or rarely in groups, brown, septate, smooth, unbranched,
proliferating percurrently through the collarette, 85.5-207.0 x 3.5-5.0 pum.
Conidiogenous cells phialidic, integrated, terminal, light brown, smooth, cylindrical,
collarette distinct or not. Conidia 1-celled, hyaline, ovoid, smooth-walled, with basal
apiculus, formed at multiple loci and aggregated in slimy heads at the apex of the
phialide, 2.5-3.0 x 2.0-2.5um. «

Material examined: SP250849.

Literature consulted: Romero, 1983.

ost

Fig. 2. Chalara aurea (x 1560), the arrow indicates a phialide. Fig. 3-4. Chloridium
virescens var. virescens (x 1560); 3. slimy head of conidia (arrow); 4. phialide with
multiple loci. Fig. 5. Cladosporium cladosporioides (x 1600). Fig. 6. Cryptophiale
kakombensis (x 480).

sf

Cladosporium cladosporioides (Fres.) De Vries in Contribution to the knowledge of
the genus Cladosporium Link ex Fr., p.57, 1952 (Fig. 5).

Conidiophores erect, dark-brown, septate, smooth, branches at the apex forming
conidial chains, 72.0-426.0 x 6.0-7.5 um. Conidiogenous cells blastic, ramoconidia
cylindrical, light brown, smooth, 1-2-celled, scars at the ends, 14.0-18.5 x 4.5-5.0
um. Conidia blastic, usually 1-celled, light-brown, ellipsoidal with pointed ends,
lemon-shaped or cylindrical, smooth-walled, with prominent scars, joined in branched
chains, 3.0-7.0 x 2.0-3.5 um.

Material examined: SP250851.

Literature consulted: Domsch ef al., 1980.

Cryptophiale kakombensis Pirozynski in Can. J. Bot., 46: 1124, 1968 (Fig. 6).
Conidiophores erect or slightly curved, setiform, single, dark-brown, septate, smooth,
thick-walled, 172.0-275.0 x 6.5-10.5 um, tapering to a pointed tip; fertile zone light-
brown, at the superior part of the conidiophore but leaving the apex visible, formed by
two layers of cells resembling a plate, at both sides of the conidiophore.
Conidiogenous cells phialidic, obscured by the cell layer. Conidia phialidic, 2-celled,
hyaline, narrow and falcate, smooth-walled, aggregated in a slimy mass at one side of
the fertile zone, 25.0-30.0 x 1.0-1.5 um.

Material examined: SP250842, SP250843.

Literature consulted: Sutton et al., 1989.

Cryptophiale udagawae Pirozynski & Ichinoe in Can. J. Bot., 46: 1126, 1968 (Fig. 7).
Conidiophores_ erect or slightly curved, setiform, single, brown, septate, smooth,
thick-walled, 102.0-161.5 x 5.0-8.0 um, 1-3 pointed dichotomous branches at the
apex; fertile zone light-brown from the upper half of the conidiophore towards its first
branch, with two layers of cells resembling a plate, at both sides of the conidiophore;
enlarged cells in the basal region. Conidiogenous cells phialidic, obscured by the
shielding cell layer. Conidia phialidic, 1-2-celled, hyaline, narrow and falcate,
smooth-walled, aggregated in a slimy mass in the fertile zone, 14.0-17.0 x 1.0-1.5 um.
Material examined: SP250841.

Literature consulted: Sutton et al., 1989.

Cylindrocladium scoparium Morgan in Bot. Gaz., 17: 191, 1892 (Fig. 8-9).
Conidiophores erect, hyaline, septate, smooth, apex iregularly branched; elongate
appendages generally present, delicate, septate, hyaline, smooth and terminating in a
clavate vesicle of 13.5-20.0 x 4.0-8.0 um at the end. Conidiogenous cells phialidic,
arising from the ultimate branches of the conidiophores, 11.5-14.0 x 2.0-2.5 um.
Conidia phialidic, 1-septate, hyaline, cylindrical, smooth-walled, single on the
phialides, but aggregated in a slimy mass, 42.0-59.0 x 3.5-4.5 um.

Material examined: SP250835.

Literature consulted: Peerally, 1991; Crous & Wingfield, 1994.

378

Fig. 7. Cryptophiale udagawae (x 830). Fig. 8-9. Cylindrocladium scoparium;
8. conidiuin (x 840); 9. apical vesicle (arrow) (x 1680). Fig. 10. Idriella falcata
(x 1140), a conidial cluster (arrow). Fig. 11. Kionochaeta ramifera (x 500).

379

Dictyochaeta simplex (Hughes & Kendrick) Hol.-Jech. in Folia Geobot. Phytotax., 19:
434, 1984 (Fig. 21).

Conidiophores erect, single or in groups of 2-3, brown, scarcely septate, smooth,
unbranched, 45.0-71.0 x 2.5-4.0 bm including the phialide; setae absent.
Conidiogenous cells phialidic, integrated, terminal, light-brown, cylindrical, with a
conspicuous collarette and sympodial growth (polyphialides). Conidia phialidic, 1-
celled, hyaline, lunate, smooth-walled, a setula at both ends, 15.0-19.0 x 2.0-3.0
ium, aggregated in a slimy mass at the apex of the phialides. Setula filiform,
hyaline, 7.0-9.0 um.

Material examined: SP250852.

Literature consulted: Hughes & Kendrick, 1968 (as Codinaea simplex)

Idriella falcata (Sutton & Hodges) von Arx in Sydowia, 34: 36, 1981 (Fig. 10, 23).
Conidiophores erect or flexuous, single, few to numerous, hyaline, delicate, smooth,
loosely aggregated, irregularly branched at the apex with the terminal branches being
the conidiogenous cells. Conidiogenous cells polyblastic and sympodial, terminal,
hyaline, delicate, subulate with inconspicuous denticles at the apex, 9.5-14.0 x 1.0-2.5
um. Conidia 1-celled, hyaline, slightly falcate, smooth-walled, with a lighter central
zone, 20.0-26.0 x 1.5-2.0 um.

Material examined: SP250855.

Literature consulted: Sutton & Hodges Jr., 1976 (as Microdochium falcatum).

Kionochaeta ramifera (Matsushima) Kirk & Sutton in Trans. Br. Mycol. Soc., 85:
715, 1985 (Fig. 11, 24).

Conidiophores erect, setiform, single, brown, septate, smooth, thick-walled, 194.0-
307.0 x 8.0-11.5 ium; with 1-14 setiform lateral branches, septate, brown and smooth,
arising approximately at the mid part of the conidiophore, from where the fertile zone
also diverges. Conidiogenous cells phialidic, lageniform, light-brown, smooth, densily
aggregated at the fertile zone. Conidia 1-celled, hyaline, falcate, smooth-walled,
aggregated in a slimy mass in the fertile zone, 7.0-8.0 x 0.5-1.0 um.

Material examined: SP250838, SP250839, SP250840.

Literature consulted: Matsushima, 1971 (as Chaetopsina ramifera); Kuthubutheen &
Nawawi, 1988.

Lauriomyces heliocephala (V. Rao & de Hoog) Cast. & Kendr. in Univ. Waterloo
Biol. Ser., 32: 26, 1990 (Fig. 12, 25).

Conidiophores erect or recurved, single or in small groups, brown, septate, smooth,
with a complex sporogenous apparatus at the apex, 62.5-98.5 (-187.5) x 4.0-6.0 um at
the base; some clusters of conidiophores associated with long filaments resembling
setae. Conidiogenous apparatus hyaline, formed by conidial chains, in dry spherical
heads. Conidia 1-celled, hyaline, cylindrical, smooth-walled, 4.5-5.0 x 1.0-1.5 yum.
Material examined: SP250854.

Literature consulted: Rao & Hoog, 1986 (as Haplographium heliocephalum).

26
Es
Cie os
mS .o
m iy
3
ire
re
2%
apd
5 Yi aaa
8 .2
Q
2%
2 §
EN
= 2
ie)
os ne
nas
ee
a S
w= oS
ms
mA
oe.
ee
=.
St
82
3
SES
Q9 9
S Bx
so)
vo .4
wm ~—
% & Ss
iS) q
eae
[a0
See
3% xy
See
ee
NS 's
A aN
nS
; ~~
i, Sea,

381

Mycoleptodiscus brasiliensis Sutton & Hodges Jr. in Nova Hedwigia, 27: 694, 1976
(Fig. 13-14).

Conidiophores slightly differentiated or indistinct. Only two small specimens were
observed, with superficial dark-brown sporodochia, spherical, one-celled width, 20.0-
24.5 tum diameter. Conidiogenous cells phialidic, ampulliform or doliiform, dark-
brown, strongly compacted, thick-walled, 6.0-8.0 x 4.0-5.5 [um, circular apical
opening, conspicuous collarette. Conidia not observed.

Material examined: SP250844.

Literature consulted: Sutton & Alcom, 1990.

Phaeoisaria clematidis (Fck1.) Hughes in Can. J. Bot., 36: 795, 1958 (Fig. 15, 26).
Conidiomata synnematous, erect, light-brown or yellowish, sometimes branched,
262.5-791.5 x 23.0-52.5 um; fertile zone approximately 2/3 of the synnema from the
apex. Conidiophores adpressed, densily compact at the base tapering towards the
apex. Conidiogenous cells oblong, light-brown to hyaline, sinuate, denticulate,
exhibiting sympodial growth. Conidia 1-celled, hyaline, ellipsoidal to obovoid,
smooth-walled, 4.0-5.5 x 1.0-1.5 um.

Material examined: SP250845.

Literature consulted: Deighton, 1974; Hoog & Papendorf, 1976.

Phaeoisaria infrafertilis Sutton & Hodges Jr. in Nova Hedwigia, 27: 219, 1976
(Fig. 16, 27).

Conidiomata synnematous, erect, dark-brown, unbranched, 250.0-500.0 x 12.0-19.5
um; fertile zone limited to the central part of the synnema, white coloured.
Conidiophores adpressed, septate and slender, densily compact at the base tapering
towards the apex. Conidiogenous cells divergent, light-brown, denticulate, exhibiting
sympodial growth. Conidia 1-celled, hyaline, falcate, smooth-walled, acute at the
base, 14.0-19.5 x 0.5-1.0 um.

Material examined: SP250846, SP250847, SP250848.

Literature consulted: Kirk, 1985.

Subulispora procurvata Tubaki in Trans. Mycol. Soc. Japan, 12: 20, 1971 (Fig. 17-
18). :

Conidiophores_ erect, straight or slightly curved, single, brown, septate, smooth,
unbranched, 52.5-106.5 x 3.5-4.5 um; basal cell lobate, apex with lighter fertile zone.
Conidiogenous cells polyblastic, integrated, apical, zigzag-like, sympodial growth,
thin scars present. Conidia 3-septate, hyaline, subuliform, smooth-walled, truncate at
the base, 21.0-26.0 x 2.0-3.0 um, with apical appendage. Appendages filiform and
hyaline, frequently forming an angle of 45-135° in relation to the main axis of the
conidia, 9.0-17.0 tum long.

Material examined: SP250836.

Literature consulted: Matsushima, 1971.

382

<<

Fig. 17-18. Subulispora procurvata,; 17. conidium (x 1870); 18. conidiophore
(x 980). Fig. 19-20. Subulispora rectilineata (x 1870); 19. conidium; 20.

conidiophore. Fig. 21. Dictyochaeta simplex (x 1560), the arrow indicates the
collarette. Fig. 22. Thozetella cristata (x 390).

383

Zs 24

25 26

2, 28 29

Fig. 23. Idriella falcata: conidia. Fig. 24. Kionochaeta ramifera: conidia. Fig. 25.
Lauriomyces heliocephala: conidia. Fig. 26. Phaeoisaria clematidis: conidia. Fig.
27. Phaeoisaria infrafertilis: conidia. Fig. 28. Subulispora rectilineata: conidia.
Fig. 29. Thozetella cristata: conidia. Scale bars = 10 um.

384

Subulispora rectilineata Tubaki in Trans.Mycol.Soc.Japan, 12:21,1971(Fig.19-20, 28).
Conidiophores erect, straight or slightly curved, single, brown, septate, smooth,
unbranched, 26.0-73.0 x 3.0-5.0 um; basal cell lobate, fertile zone at the apex.
Conidiogenous cells polyblastic, integrated, terminal, zigzag-like, sympodial growth,
typical scars present. Conidia 1-2-septate, hyaline, subuliform, smooth-walled,
truncate at the base, tapering at the apex, 35.0-41.5 x 1.5-2.0 um.

Material examined: S$P250837.

Literature consulted: Hoog, 1985.

Thozetella cristata Pirozynski & Hodges Jr. in Can. J. Bot., 51:168,1973 (Fig. 22, 29).
Conidiomata synnematous, erect, light-brown, transverse growth zone present or not,
64.5-110.5 x 38.0-54.0 um. Conidiophores densily compact, hyaline when separated,
septate, with one phialide at the apex. Conidiogenous cells phialidic, hyaline,
cylindrical or bacilliform, smooth. Conidia 1-celled, hyaline, lunate, smooth-walled,
hyaline setule at the ends, 9.0-14.5 x 1.0-2.0 uum. Microawns falcate or sigmoid,
hyaline, smooth-walled and refracting at the fertile zone.

Material examined: SP250850.

Literature consulted: Pirozynski & Hodges Jr., 1973.

CONCLUSIONS

The study of filamentous fungi at the ERJI is in a pioneering stage. The
results give important information about the geographical distribution of the following
taxa that are mentioned for the first time in Brazil: Lauriomyces heliocephala and
Subulispora rectilineata. This paper represents an identification aid for other
researchers in this wild tropical area and contributes to the knowledge on its fungal
biodiversity.

ACKNOWLEDGEMENTS

The authors are deeply indebted to Mr. Marcos G. Sobral (Universidade
Federal do Rio Grande do Sul) for his help on collecting the Alchornea triplinervia
leaves. We also thank Mrs. Cristina Y. Umino and Miss Lilian D. Martins (Fundagao
Tropical “André Tosello”) for assistance with the photographs and the drawings,
respectively, and Dr. Keith Seifert (Agriculture and Agri-Food, Canada) for revising
this manuscript.

LITERATURE CITED

ARX, J.A. von. 1981. Notes on Microdochium and Idriella. Sydowia, 34: 30-38.

ATTILI, D.S. 1994. Isolamento, identificagao e ecologia de fungos celuloliticos do
solo da Estacgao Ecolégica de Juréia-Itatins, SP. Rio Claro, UNESP, 148 p. Tese
(Doutorado) - Instituto de Biociéncias, Universidade Estadual Paulista.

ATTILI, D.S., ZAVATINI, J.A., PINTO, I.M.A. 1995. Caracteristicas climaticas da
Estagao Ecolégica de Juréia-Itatins, Mata Atlantica, Sao Paulo, Brasil. Geografia,
20 (1), in press.

385

CASTANEDA RUIZ, R.F., KENDRICK, B. 1990. Conidial fungi from Cuba: I. Univ.
Waterloo Biol. Ser., 32: 1-53.

CHATANON, L., KIGAWA, R., KAWASAKI, H., ATTILI, D.S., CANHOS, V.P.,
KURAISHI, H., SUGIYAMA, J. 1994. The diversity of soil microfungi in the
Atlantic Rain Forest of Brazil: a preliminary taxonomic analysis. In: Annual
Reports of International Center of Cooperative Research in Biotechnology. Japan,
Vals.

COOKE, R.C., RAYNER, A.D.M. 1984. Ecology of saprotrophic fungi. Longman,
New York. 415 p.

CORTESAO, J. 1989. Juréia. Ed. Index, Sao Paulo. 133 p.

CROUS, P.W., WINGFIELD, M.J. 1994, A monograph of Cylindrocladium, including
anamorphs of Calonectria. Mycotaxon, 51: 341-435.

DE VRIES, G.A. 1952. Contribution to the knowledge of the genus Cladosporium
Link ex Fr. Uitgeverij & Drukkerij Hollandia, Baarn. 121 p.

DEIGHTON, F.C. 1974. Four synnematous Hyphomycetes. Trans. Br. mycol. Soc., 62
(2): 243-252.

DOMSCH, K.H., GAMS, W., ANDERSON, T.-H. 1980. Compendium of soil fungi.
Academic Press, London, V. 1. 859 p.

GAMS, W. , HOLUBOVA-JECHOVA, V. 1976. Chloridium and some other
dematiaceous hyphomycetes growing on decaying wood. Stud. Mycol., 13: 1-99.
GARLIPP, A.B. 1995. Isolamento e identificagao de fungos filamentosos do solo do
Banhado Grande, na Estacao Ecolégica de Juréia-Itatins, SP. Rio Claro,
UNESP, 94 p. Dissertagio (Mestrado) - Instituto de Biociéncias, Universidade

Estadual Paulista.

HARLEY, J.L., WAID, J.S. 1955. A method of studying active mycelia on living roots
and other surfaces in the soil. Trans. Br. mycol. Soc., 38: 104-118.

HOLUBOVA-JECHOVA, V. 1984. Lignicolous Hyphomycetes from Czechoslovakia
7. Chalara, Exochalara, Fusichalara and Dictyochaeta. Folia Geobot.
Phytotaxon. , 19: 387-438.

HOOG, G.S. 1985. Taxonomy of the Dactylaria complex. IV. Dactylaria, Neta,
Subulispora and Scolecobasidium. Stud. Mycol., 26: 1-60.

HOOG, G.S., PAPENDORF, M.C. 1976. The genus Phaeoisaria. Persoonia, 8 (4):
407-414.

HUGHES, S.J. 1958. Revisiones hyphomycetum aliquot cum appendice de nominibus
rejiciendis. Can. J. Bot. , 36: 727-836.

HUGHES, S.J.,. KENDRICK, W.B. 1968. New Zealand fungi. 12. Menispora,
Codinaea, Menisporopsis. N.Z.J. Bot., 6: 323-375.

KIRK, P.M. 1985. New or interesting microfungi. XIV. Dematiaceous Hyphomycetes
from Mt. Kenya. Mycotaxon, 23: 305-352.

KIRK, P.M., SUTTON, B.C. 1985. A reassessment of the anamorph genus
Chaetopsina (Hyphomycetes). Trans. Br. mycol. Soc., 85 (4): 709-717.

KUTHUBUTHEEN, A.J., NAWAWI, A. 1988. Two new species of Kionochaeta
(Hyphomycetes) and K. ramifera from Malaysia. Trans. Br. mycol. Soc., 90 (3):
437-444.

MATSUSHIMA, T. 1971. Microfungi of the Solomon Islands and Papua-New Guinea.
Nippon Printing, Osaka. 78 p. (published by the author).

386

MORGAN, A.P. 1892. Two new genera of Hyphomycetes. Bot. Gaz., 17: 190-192.

NAG RAJ, T.R., HUGHES, S.J. 1975. A monograph of Chalara and allied genera.
Wilfr. Laurier Univ. Press, Waterloo. 200 p.

PEERALLY, A. 1991. The classification and phytopathology of Cylindrocladium
species. Mycotaxon, 40: 323-366.

PIROZYNSKI, K.A. 1968. Cryptophiale, a new genus of Hyphomycetes. Can. J. Bot.,
46: 1123-1127.

PIROZYNSKI, K.A., HODGES JR., C.S. 1973. New Hyphomycetes from South
Carolina. Can. J. Bot.,51: 157-173.

RAO, V. , HOOG, G.S. 1986. New or critical Hyphomycetes from India. Stud. Mycol.,
28: 1-84.

ROMERO, A.I. 1983. Contribucion al estudio de los hongos xilofilos de la Argentina.
I. Deuteromycotina en Eucalyptus viminalis (Myrtaceae). Bol. Soc. Argent. Bot.,
22 (1-4): 57-79.

SUTTON, B.C., ALCORN, J.L. 1990. New species of | Mycoleptodiscus
(Hyphomycetes). Mycol. Res., 94: 564-566.

SUTTON, B.C., HODGES Jr., C.S. 1976. Eucalyptus microfungi: Microdochium and
Phaeoisaria species from Brazil. Nova Hedwigia: 27: 215-222.

SUTTON, B.C., HODGES Jr., C.S. 1976. Eucalyptus microfungi: Mycoleptodiscus
species and Pseudotracylla gen. nov. Nova Hedwigia: 27: 693-700.

SUTTON, B.C., NAWAWI, A., KUTHUBUTHEEN, A.J. 1989. Additions to
Belemnospora and Cryptophiale from Malaysia. Mycol. Res., 92(3): 354-358.

TRAPPE, J.M., SCHENCK, N.C. 1982. Taxonomy of the fungi forming
endomycorrhizae. In: Schenck, N.C. (ed.). Methods and principles of
mycorrhizae research. St. Paul: The American Phytopathological Society. p.1-9.

TUBAKI, K., YOKOYAMA, T. 1971. Notes on the japanese Hyphomycetes V. Trans.
Mycol. Soc. Japan, 12: 18-28.

MYCOTAXON

Volume LX, pp. 387-395 October-December 1996

BRIDGEOPORUS, A NEW GENUS TO ACCOMMODATE
OXYPORUS NOBILISSIMUS
(BASIDIOMYCOTINA, POLYPORACEAE)

Harold H. Burdsall, Jr., Thomas J. Volk,
Center for Forest Mycology Research,
Forest Products Laboratory *, Forest Service, United States Department of
Agriculture, One Gifford Pinchot Dr., Madison, WI 53705 USA

& Joseph F. Ammirati, Jr.
Department of Botany, KB-15, University of Washington,
Seattle WA 98195 USA

ABSTRACT: The new genus Bridgeoporus is proposed to accommodate
Oxyporus nobilissimus W.B. Cooke. Bridgeoporus is associated with a
brown rot of wood, as opposed to the superficially similar genera
Oxyporus and Rigidoporus, which are associated with white rot.
Bridgeoporus lacks clamp connections at the septa, is monomitic,
possesses pseudocystidia, and has a unique configuration of fascicles of
hyphae making up the upper surface of its pileus. This combination of
characters is found in no previously described genus.

* The Forest Products Lab is maintained in cooperation with the
University of Wisconsin-Madison. This publication was written and
prepared by U.S. government employees on official time, and it is
therefore in the public domain and not subject to copyright.

INTRODUCTION

Oxyporus nobilissimus W.B. Cooke (Cooke, 1949) has attracted a
great deal of attention in the past several years (Christy, 1991; Coombs,
1991) because of its large size, uncommon occurrence, unique appearance,
and association with very large host trees. Commonly called the "Fuzzy
Sandozi," it has been reported infrequently (WTU Herbarium records;
Trappe, 1990; Christy, 1991) since its discovery in 1943. Currently, living

388

specimens are known from only six sites in Oregon and Washington. The
fungus produces perennial basidiomata that may weigh up to up to 130 kg
and are usually associated with large diameter (at least 1 m) Abies procera
Rehd., but occasionally with equally large Abies amabilis (Dougl.) Forb.
or possibly Tsuga heterophylla (Raf.) Sarg. It is the first fungus to be
listed as an endangered species by any private or public agency in the
United States, having been listed as such by the Oregon Natural Heritage
program (Christy, Pers. Comm.; Lizon, 1995). O. nobilissimus has also
been listed by the Forest Ecosystem management team (FEMAT; anon.,
1994) as one of 252 species of fungi that must be surveyed for in the
habitat of the northern spotted owl.

The genus Oxyporus (Bourd. & Galz.) Donk accommodates
species with a monomitic hyphal system, simple-septate hyphae, and
causing white rot. Oxyporus nobilissimus has been reported to cause a
brown rot (Cooke, 1949), but the species has been left in Oxyporus (e.g.,
by Gilbertson & Ryvarden, 1987) because of a lack of specific studies
demonstrating the type of rot. It has also been placed in Fomes (Fr.) Fr.
(Lowe, 1955). However, because of the following combination of
characteristics, 1.e. simple-septate hyphae, association with brown rot,
pseudocystidia of tramal origin, and closely-appressed hyphae in fascicles
making up the upper surface of its pileus, it is being placed in a new
genus, Bridgeoporus.

METHODS AND MATERIALS

Relatively small (5-6 cm) portions of basidiomata were carefully
excised from living specimens and cultured according to Gilbertson &
Ryvarden (1986) on 1.5% malt extract 2% agar, plus 2 mg/l benomyl! and
100 mg/l streptomycin. Microscopic characters of the basidiomata were
determined by mounting thin sections in 3% KOH plus 1% phloxine or in
Melzer’s reagent (Gilbertson & Ryvarden, 1986). Line drawings were
made with the aid of a camera lucida and drawing tube. Putative cultures
and portions of basidiomata have been preserved at CFMR, Madison,
Wisconsin. Herbarium abbreviations are as in Holmgren et al. (1990).
Colors are as in Ridgway (1912). PCR amplification and sequencing was
performed according to Nakasone & Sytsma (1993).

389
RESULTS AND DISCUSSION

Although collected basidiospores failed to germinate, we collected a
number of putative cultures from tissues of four basidiomata and from the
brown-rotted wood. We amplified and sequenced the internal transcribed
spacer (ITS) region of the ribosomal DNA repeat from DNA obtained
from basidiospore drops and from putative cultures; the DNA sequence
from the spore drops (two samples) and 3 putative cultures did not match.
Thus we could not positively conclude that any of the tissue cultures were
that of Oxyporus nobilissimus. However, O.nobilissimus has been
reported to cause a brown rot because of the decay observed in the wood
attached to the type specimen (Cooke, 1949). Our observations of wood in
association with living specimens (whenever possible) also indicate brown
rot. We are confident that O.nobilissimus causes a brown rot. The brown
rot associated with the basidiomata is in contrast to that of the type species
of Oxyporus, Polyporus connatus Weinm. [ = Polyporus populinus
Schum.:Fr., = Oxyporus populinus (Schum.:Fr.) Donk ], which causes a
white rot.

Besides the difference in rot characteristics, which are considered
important at the generic level (e.g. Nobles, 1958; Donk, 1960; Ryvarden,
1991), Oxyporus (Bourd. & Galz.) Donk differs from O.nobilissimus in
having true cystidia that arise from the subhymenium rather than
pseudocystidia, which are differentiated hyphal end cells that arise in the
trama. Species of Rigidoporus Murr. are similar to O.nobilissimus in
possessing pseudocystidia, similar basidiospore and basidium shape and
size, and both thin- and thick-walled generative hyphae, but it causes a
white rot. Rigidoporus is probably the closest genus for placement of this
species, but its members cause a white rot and lack the unique pileus
surface hyphal structure of O.nobilissimus. Lowe (1955) placed
O.nobilissimus in Fomes (Fr.) Fr. at a time when that genus encompassed
a wide variety of large, conk-shaped species. It is no longer an
appropriate placement since the concept of Fomes has been narrowed to
include only trimitic species, with clamped generative hyphae and sclerids,
and causing a white rot.

Other genera with simple-septate hyphae that we considered for
placement of this species included Physisporinus Karst., but its members
have resupinate, soft, waxy basidiomata and are associated with a white
rot. Pycnoporellus Murr., em. Kotl. & Pouz., contains species that cause
a brown rot and form bright orange to rust colored basidiomata that turn
deep red in KOH. Postia Fr. (= Oligoporus Bref.) contains species whose
members have clamp-connections on generative hyphae. In addition, the

390

broadly ovoid basidiospore shape in O.nobilissimus is very different from
known Postia species. None of the above genera contain species with the
unique hyphal structure of hyphae on the pileus surface that is found in
O.nobilissimus. Since no described genus is appropriate, we propose a
new genus, Bridgeoporus, to accommodate this unique species.

Bridgeoporus Volk, Burdsall & Ammirati gen. noy. Figs. 1-8.
Basidiomatibus perennis, effusis-reflexis vel substipitatis, imbricatis,
concrestis, suberosis, lignosis, pileis dense strigosis, fibris ramosis
hispidis, zonatis, systemate hyphae monomitico, hyphis hyalinis, efibulatis,;
pseudocystidiis hyalinis; basidiosporis hyalinis, laevibus, nonamyloideis.
Consociateus una cum tabes brunneo.

Etymology: Bridgeoporus m. Named for William Bridge Cooke (widely
known as "Bridge"), who described Oxyporus nobilissimus.

Type species: Bridgeoporus nobilissimus (W.B. Cooke) Volk, Burdsall, &
Ammirati comb. nov.
basionym: Oxyporus nobilissimus W.B. Cooke, Mycologia 41: 444 (1949)
= Fomes nobilissimus (W.B. Cooke) Lowe, Mycologia 47:219
(1955)

Basidiomata perennial, 30-140 x 25-95 x 30-100 cm, sessile, ungulate,
imbricate, or centrally substipitate, with the form depending on location on
host tree (snag, stump, or living) (Fig. 1, 2) often occluding small twigs
and other objects . Texture fibrous, rubbery and tough when fresh. Pileus
surface (Fig. 3) a dense mat of white mycelial fibers in youth, often
somewhat agglutinated at the tips, becoming cinnamon brown or darker in
age and usually reaching several mm in length, often appearing green due
to epiphytic association with several species of unicellular algae. Context
up to 1.5 cm thick, white, tough, rubbery, and fibrous when fresh,
cinnamon-buff to ochraceous, hard and brittle when dried. Pores
concolorous with context, round, 2 per mm, 2-7 mm long in mature
layers, not becoming stuffed, stratified, with a layer of sterile tissue 2-3
mm thick between successive pore layers.

Figs. 1-3. (opposite page) Macroscopic characters of Bridgeoporus nobilissimus.
1. Sessile form growing on the side of a snag. Note brown rotted wood behind
the somewhat inconspicuous specimen in the center. 2. Centrally stipitate form
growing on the ground from roots of a snag. 3. Closeup of pileus surface with
coarse hairs. For information on how to obtain color computer images of these
and other photographs of B.nobilissimus please contact T.Volk, whose current e-
mail address is <tjvolk@facstaff.wisc.edu >

pee.

Fibers of the pileus surface 10-30 mm X (50)-60-75 um, with frequent
branching and anastomosing, showing layering as evidence of seasonally
added increments of growth, new growth of the fibers occurring in
discrete tufts of agglutinated hyphae scattered over the pileus surface,
especially near the margin. The fibers are composed of bundles of simple-
septate, parallel hyphae, 2-3 4m diam, hyaline to yellow-brown, simple-
septate, thin- to slightly thick-walled, infrequently branching.

Context hyphae 3-4 wm diameter, thin-walled to thick-walled, simple- -
septate, hyaline to pale yellow-brown, smooth, rarely branched. Tramal
hyphae like those of context, but some becoming thick walled and growing
intrusively through the subhymenium, becoming pseudocystidia.
Pseudocystidia up to 125 um x 6-12 um, cylindrical to broadly subulate,
arising deep in the tramal tissue, evident before formation of basidia, walls
slightly thickened or up to 4 um thick in age, often with a hyaline,
crystalline cap. Basidia 12-18 x 4-10 wm, pyriforme, 4-spored, simple-
septate at base. Sterigmata 2-3 ym long. Basidiospores 5.5-6.5 x 3.5-4.5
um, broadly ovoid, hyaline, smooth, thin-walled, negative in Melzer’s
reagent.

Habitat: Occurring singly or occasionally imbricate on old growth Abies
procera (1-2 m or more diameter at breast height), rarely Abies amabilis
or possibly Tsuga heterophylla. Position on the host is variable. It has
been found near the root collar of living trees or snags, on sides of snags
not far from the ground (1 m maximum height), and on old cut stumps,
but not known to occur on downed logs.

Distribution: Known only from the Cascade Mountain Range in
Washington and Oregon and the Coast Range on the Olympic Peninsula in
Washington. Locations where basidiomata are found were given letter
designations (e.g. AC, GM, LM, SP, HT, NR) based on their location.
Number in parentheses following the location indicate number of
basidiomata found at that site. Since these are perennial basidiomata of
rarely-found fungi that are still extant, we decline to give exact locations
in order to protect them.

Specimens examined: A.H. Smith No. 31102, holotype MICH, isotype WTU, Mt. Rainier
National Park, WA, Abies sp., 1948; Extant basidiomata (portions of which are at CFMR):,
AC (3), King Co., Mt. Baker-Snoqualmie National Forest, WA, A.procera, 1992; GM (4)
Cowlitz Co., Mt. St. Helens National Monument, WA, A.procera, 1992; LM (6) Multnomah
Co., Mt. Hood National Forest, OR, A.procera, T.heterophylla, 1992, 1994; SP (15) Linn
Co., Bureau of Land Management, OR, A.procera, 1991, 1992; HT (2) Olympic National
Forest, Gray’s Harbor Co., WA, Abies amabilis, 1992, 1993; NR (2) Mt. Rainier National
Park, Pierce Co., WA, A.procera, 1992.

393

rus nobilissimus. Fig. 4. Context
8. Basidiospores.

of Bridgeopo
menium. 7. Basidia.

>
WN

5
eS
eo
-
Qs
eaS
=
QS
Ooo
9 6
Qo
fe)

5 5
nth
=m
wo
si

va
Bp Qe
=~ Py
mB

394

REMARKS: The type specimen was reported on Tsuga heterophylla (Cooke, 1949).
However, microscopic examination of the wood attached to the base of the isotype is
that of an Abies sp. (Regis Miller, Center for Wood Anatomy Research, Forest
Products Lab, Personal Communication). This is consistent with the vast majority of
specimen/host associations noted for B. nobilissimus. However one of the specimens
in Multnomah County has been found at the base of a large T. heterophylla snag next
to a large Abies procera snag. The host association of this specimen is unclear.

Cooke (1949) considered the fungus to have a dimitic hyphal system. Although
thin-walled and thick-walled hyphae are present in the basidiomata, the latter should
not be considered skeletal hyphae because of their abundant septa, but rather as
sclerified generative hyphae. We concur with Gilbertson & Ryvarden (1987) who
report O. nobilissimus as monomitic. Additional illustrations and descriptions of
micro- and macroscopic characteristics can be found in Lowe (1955, 1957),
Gilbertson & Ryvarden (1987), Coombs (1991) and Christy (1991).

The mat of mycelia on the upper surface is not solid, but very porous, containing
cracks and crevasses that allow the accumulation of water and organic debris.
Unicellular algae, including Coccomyxa sp. and Charicium spp., also occur. Our
studies indicate that the algae appear to be living among fungal hyphae and not
endophytically. In addition, some vascular plants (e.g. Oxalis sp. and pteridophytes)
and bryophytes sometimes occur in and on the surface of the pileus.

As indicated in the description, a layer of external hyphae covers the previous
year’s pore surface before the initiation of a new pore layer. Each new layer arises
from coalescence of discrete patches of mycelium formed external to the old pores.
The pores begin as indentations on the layer of confluent sterile hyphae.

ACKNOWLEDGMENTS: We thank James Trappe, as well as Don H. Coombs and
M. Maggie Rogers (the editors of Mushroom, the Journal of Wild Mushrooming)
for renewing interest in this fungus. We thank Glenn Walker, James H. Ginns, David
Shaw, Claire Hibler, Judy Roger, Lorelei Norvell, Elisabeth Eyestone, and Don and
Bonnie Grandorff for help in finding and collecting specimens and putative cultures of
this fungus. We thank Karen K. Nakasone, Rita M. Rentmeester and Kelly Collins for
testing the identity of the cultures using PCR fingerprinting and DNA sequencing. We
thank Linda and James Graham for identification of algal associates and Regis Miller
for identification of host tree species. We thank Karen K. Nakasone and Michael J.
Larsen for help with the Latin diagnosis. We thank Michael J. Larsen and Robert L.
Gilbertson for review of the manuscript. We also thank Erast Parmasto, Leif
Ryvarden, and Tuomo Niemela for helpful discussions on the proper placement of this
species.

395

LITERATURE CITED

Anonymous. 1994. Standards and Guidelines for management of habitat for late-
successional and old-growth forest related species within the range of the northern
spotted owl. Attachment A to the Record of decision for amendments to Forest
Service and Bureau of Land Management planning documents within the range of
the northern spotted owl. US Government Printing Office, Region 10.

Christy, J. 1991. The most noble polypore--- Endangered or already gone to
passenger pigeon-land. Mushroom, the Journal of Wild Mushrooming 9(3):11

Cooke, W.B. 1949. Oxyporus nobilissimus and the genus Oxyporus in North
America. Mycologia 41:442-455.

Coombs, D.H., 1991. Looking for a big fuzzy one. Mushroom, the Journal of Wild
Mushrooming 9 (4):5-8

Donk, M. 1960. The generic names proposed for Polyporaceae. Persoonia 1:173-302.

Gilbertson, R.L. and L. Ryvarden. 1986. North American Polypores. Vol. 1
Abortiporus-Lindtneria. Oslo: Fungiflora. pp. 1-436.

Gilbertson, R.L. and L. Ryvarden. 1987. North American Polypores. Vol. 2.
Megasporoporia-Wrightoporia. Oslo: Fungiflora. pp. 437-885.

_ Holmgren, P.K., N.H. Holmgren, and L.B. Barnett. 1990. Index Herbariorum.
Eighth Edition. New York Botanical Garden: 693 pp.

Lizon, P. 1995. Preserving the biodiversity of fungi. Inoculum 46 (6): 1-4.

Lowe, J.L., 1955. Perennial polypores of North America III. Fomes with context
white to rose. Mycologia 47:213-224

Lowe, J.L., 1957. Polyporaceae of North America: The genus Fomes. State
University College of Forestry at Syracuse University, Technical Publication No.
80. 97 pp.

Nakasone, K.K. and K.J. Sytsma. 1993. Biosystematic studies on Phlebia acerina,
P.rufa, and P.radiata in North America. Mycologia 85:996-1016.

Nobles, M.K. 1958. Cultural characters as a guide to the taxonomy and phylogeny of
the Polyporaceae. Canad. J. Bot. 36:883-926

Ridgway, R. 1912. Color Standards and Color Nomenclature. 53 plates.

Ryvarden L. 1991. Genera of Polypores: Nomenclature and Taxonomy. Fungiflora,
Oslo, Norway. Synopsis Fungorum 5. 363 pp.

Trappe, J.M. 1990. The "most noble" polypore endangered. Jn Ancient Forests of
the Pacific Northwest. Elliot A Norse. The Wilderness Society. Island press.
Washington DC pp. 126-127.

MYCOTAXON

Volume LX, pp. 397-414 October-December 1996

A CHECK LIST OF HIGHER MARINE FUNGI ON
WOOD FROM DANISH COASTS

J. KOCH and K.R.L. PETERSEN

Department of Plant Biology, The Royal Veterinary and Agricultural University,
Thorvaldsensvej 40, DK-1871 Frederiksberg C, Denmark

Abstract

72 higher marine fungi on wood from the Danish coast are presented including 15
species as new records for Denmark. The fungi are listed from 12 localities on the west
coast of Jutland and from 35 localities on the coasts of inner Danish waters. Comments
are given on selected species. The list is based on 988 samples which are kept at the
herbaria C and CP, Copenhagen.

Introduction

Generally, information on the occurrence of fungal species in a country or a greater
area is difficult to obtain. As a rule, troublesome studies of literature and studies of
material deposited in different herbaria are required. To remedy this the following
check list for higher marine and arenicolous fungi on wood along the Danish coasts was
prepared.

Materials and methods

The fungi are from driftwood deposited on beaches, from wood caught between stones
functioning for coastal protection, and from stationary wood such as harbour timber.
Some of the material was incubated with moist sterilized sand in plastic boxes for
several months. The fungi are listed in table 2.

The material is divided into two groups, one from the west coast of Jutland and one
from the inner Danish coast. This is due to the difference in salinity of the coastal
waters in the two areas - at the west coast of Jutland the salinity is close to 3,1% and
at the inner Danish coast between 1 and 2% (Hermann, 1968). Each locality has been
given a number referring to the list of localities, table 1, and to the map, fig. 1.

In table 2 the quoted literature is referring to earlier published collections from
Denmark. Where no literature is indicated the species is a new record for Denmark.

398

All collections mentioned have been studied by the authors. The study has been
substantially faciliated by the keys prepared by Kohlmeyer & Volkmann-Kohlmeyer
(1991).

The material, which consists of 988 samples, is kept at the herbarium (CP), The Royal
Veterinary and Agricultural University, Copenhagen, and the Botanical Museum (C),
University of Copenhagen.

Results and discussion

To some degree the number of collections give an impression of the frequency of
occurrence of single species. However, a strict quantification cannot be read from table
2 as the collecting technique has not been adapted for this purpose. A grouping taking
into account the number of collections of a species could roughly divide the fungi with
regard to frequency into the following three groups, where the species in each group
are arranged according to their frequency.

Frequency 1 (128-40 collections)

Monodictys pelagica, Arenariomyces trifurcatus, Nereiospora comata, Corollospora
maritima, Halosphaeria appendiculata, Remispora Stellata, R. maritima, Ceriosporopsis
halima, Halosphaeriopsis mediosetigera, Ceriosporopsis tubulifera, Amylocarpus
encephaloides, Lulworthia fucicola, Dictyosporium pelagicum, Zalerion maritimum,
Dryosphaera navigans, Ocostaspora apilongissima, Marinospora calyptrata, Lulworthia
lignoarenaria, Trichocladium alopallonella. - 19 species.

Frequency 2 (39-10 collections)

Remispora pilleata, Kohlmeyeriella tubulata, Nereiospora cristata, Carbosphaerella
leptosphaerioides, Kirschsteinothelia maritima, Remispora galerita, Haligena
elaterophora, Leptosphaeria oraemaris, Corollospora lacera, Digitatispora marina,
Appendichordella amicta, Sphaerulina oraemaris, Cirrenalia macrocephala,
Ceriosporopsis caduca, Groenhiella bivestia, Leptosphaeria pelagica, Carbosphaerella
pleosporoides, Haligena salina, Anisostagma rotundatum, Limacospora sundica,
Zalerion varium, Nia vibrissa, Orbimyces spectabilis, Marinospora longissima,
Torpedospora ambispinosa. - 25 species.

Frequency 3 (9-1 collections)

Eiona tunicata, Halosarpheia viscosa, Marinosphaera mangrovei, Marisolaris ansata,
Ondiniella torquata, Dictyosporium toruloides, Massarina thalassiae, Savoryella
— lignicola, Cumulospora marina, Halosarpheia trullifera, Nais inornata, Capronia
ciliomaris, Biflua physasca, Bovicornua intricata, Nautosphaeria cristaminuta (fig. 10),
Asteromyces cruciatus, Trichocladium achrasporum, Biconiosporella corniculata,
Diplodia oraemaris, Remispora quadriremis, Lautisporopsis circumvestita (fig. 11),
Chaetomastia typhicola, Chaetosphaeria chaetosa, Corollospora luteola (+ Sigmoidea
luteola), Lindra inflata, Remispora crispa, R. spinibarbata, Trichocladium constrictum.
- 28 species.

The general impression is that species with frequency 1 are common, which is in close
agreement with findings by Schmidt (1974) at the coast of Rigen in the Baltic sea and
by Schaumann (1975) in the Weser-Estuary and Helgoland at the North sea coast.

399

Species with frequency 3 are rare or very rare.

Most of the wood samples from the west coast of Jutland were collected as driftwood
on the beaches. Together with the special beach characters (Koch, 1974) at the west
coast, and the salinity of the coastal water, this could explain the prevalence of the
arenicolous species Corollospora lacera, Kohlmeyeriella tubulata and Nereiospora
comata in this area, and corresponding to that the general absence of the obligate
marine species Halosphaeria appendiculata, Lulworthia fucicola and Marinospora
calyptrata.

Schaumann (1975) states that "Ascomycetes dominate in the more saline habitats with
the Deuteromycetes predominantly in the lower saline and more limnetic parts of the
estuary". Concerning the Deuteromycetes, this also seems to be the case, when
comparing material from the west coast of Jutland with material from the inner Danish
coasts. Especially dominant here are Cirrenalia macrocephala, Dictyosporium
pelagicum, Monodictys pelagica and Trichocladium alopallonella. The same trend was
found by Schmidt (1974).

The following 15 species are new records for Denmark: Biconiosporella corniculata,
Capronia ciliomaris, Chaetomastia typhicola, Corollospora luteola (+ Sigmoidea
luteola), Haligena salina, Leptosphaeria pelagica, Marinosphaera mangrovei,
Marinospora longissima, Massarina thalassiae, Ondiniella torquata (fig. 9), Remispora
crispa, Sphaerulina oraemaris, Torpedospora ambispinosa, Cumulospora marina,
Trichocladium constrictum.

Comments on single species

Capronia ciliomaris. This species is considered exclusively as growing on bark (fig.
3), but one collection here is from heavily decayed wood. We have not been able to
observe the subterminal, radiating filamentous appendages of the ascospores. We have
only seen mucus fibrils surrounding the spores (fig. 4). This also apply to spores from
the type material (J. Kohlmeyer 226, 253 in Herb. Mus. Bot. Berol.). In all the
aspects, the type material conforms with our collections.

Ceriosporopsis caduca. By Koch (1974) wrongly cited sub nomen C. longissima
Kohlm. and by Koch & Jones (1983) sub nomen C. cambrensis I.M. Wilson. In the
Danish material of C. caduca, we have not observed the shedding of the outer sheath
from the polar appendages as mentioned by Jones & Zainal (1988), but catenophyses
and filamentous catenophyses are abundant.

Chaetomastia typhicola. We record this species with hesitation since it has been
described from Typha, Juncus and Phragmites and not from wood. Microscopic
characters are in agreement with Kohlmeyer & Kohlmeyer (1979) who consider the
species a facultative marine fungus.

Dictyosporium toruloides. This species is considered a facultative marine fungus. We
have found it on intertidal wood and on a mooring post of oak in Svanemellen Harbour
(39) 60 cm below daily water. Ellis (1971) wrote "Common on wood and herbaceous
stems" indicating it as a terrestrial species.

Dryosphaera navigans. Early records (Farrant et al. 1985, Koch, 1974, 1985, and
Koch & Jones, 1983) of this species were given sub nomen Crinigera maritima I.
Schmidt (see Koch & Jones, 1989).

400

Halosarpheia viscosa. In the quoted collections the ascospores are mostly separated
from the ascomata in masses without showing asci, but when seen, they are thinwalled.
The ascospore dimensions are also close to Ceriosporopsis hamata Hohnk and
Halosarpheia retorquens Shearer et Crane. We have therefore not been able to assign
our collections with confidence to any of the mentioned taxa. We consider the
collections as H. viscosa, sensu lato.

Also we have 8 collections with Halosarpheia spores (thinwalled), showing persistent
Aniptodera-like asci. We have found no genus to accomodate this species.
Marinosphaera mangrovei. Since this species is mainly reported from tropical zones,
we have compared (LM) our collections with the type material. This was found to
conform with our fungus. Ascospores from a Danish collection (figs 5-7) showed, that
they were covered by a fibrillar material (fig. 2) as shown by Read et al. (1995).
Marinospora longissima. See comments relating to Ceriosporopsis caduca.
Massarina_thalassiae. This species is common on oak in the intertidal zone at
Strandegaard Dyrehave (44), but has not been found on oak or other wood substrates
in similar positions from any Danish localities. The species is described by Kohlmeyer
& Volkmann-Kohlmeyer (1987) on different wood substrates from the tropics. With
regard to ascospore dimension the Danish collections are very close (30-36,4-40 x 10-
11,9-12 wm, n=31) to dimension given by Kohlmeyer & Volkmann-Kohlmeyer (1987)
(31,0-37,.8-43 ,5(-47,1) x 9,7-12,2-14,1 um, n=215). Asci measured in situ conform
with measurements by Kohlmeyer & Volkmann-Kohlmeyer (1987). M. thalassiae has
not been recorded before in the temperate zone (fig. 8).

Ocostaspora apilongissima. In Koch (1974) as a variety of Halosphaeria appendiculata.
Remispora crispa. This is the first record of the species outside the tropical region. It
was found on a mooring post of oak in Svanemellen Harbour (39), where the harbour
bassin for many years was kept free of ice because of outlet of cooling water from an
electricity plant. Nevertheless it is with some hesitation we list this species.
Remispora galerita. In Koch (1974) sub nomen Lentescospora submarina Linder.

Table 1. List of Danish localities. 1-12 west coast of Jutland, 13-47 coast of inner
Danish waters. See fig. 1 and table 2.

Number/Locality Number/Locality Number/Locality

1. Tannishus 17. Havhuse 33. Niva

2. Grenhgj 18. Senderskov 34. Rungsted

3. Svinklov 19. Skovgaarde 35. Vedbek

4. Hjardemal 20. Helgenes 36. Skodsborg

5. Vigsa 21. Staksrode Skov 37. Tarbek

6. Hanstholm 22. Varbjerg Strand 38. Skovshoved

7. Stenbjerg 23. belo 39. Svanemollen Havn

8. Lyngby 24. Torrese 40. Saltholm

9. Fjaltring 25. Lange 41. Koge

10. Husby 26. Lerchenborg 42. Stevns Klint (Hajerup)
11. Argab 27. Hundested 43. Hojstrup |
12. Rome 28. Liseleje 44. Strandegard Dyrehave
13. Bygholm 29. Tisvildeleje 45. Lilleklint

14. Glyngere 30. Rageleje 46. Marrebek

15. Skive 31. Hornbek 47. Bote

16. Hals Fergeleje 32. Helsingor

Fig. 1. Map of Denmark showing localities for collecting lignicolous and arenicolous
marine fungi. See list of localities table 1.

x2 ‘n
a UW | a 3 ‘Y ‘3 G

1‘y =I ar ‘q

br “6€ “17 € 3 0
Ly ‘9b
‘by ‘Th ‘Ih “9E-ZE “OE
bOC, SCE. CG OC LL
Ce vi 0 oC col $6 EC

W9G-5e ChE EC
Te ST-S1-wI=S -C CC vel ¢

6¢ SI cl 0

Sp-Eb “6E “LE-SE “TZ
ESC =v oC lecanl OS VC 9¢

TLOI “ST-V ‘PT Uoaryrouorg
“YOSIOJSIIIIJI “JSUT JJOJOA - WUeU
-neYyos vIDjNIIULOD D]Ja/odso1u0jig

pS6l ‘OF :€
UdIARYIOUMIIIG YOSIOJSIIIIJAJ JSUT “JJOIDA,
- YUYOH snqwoinfisl SadKwmoiiDualy

L861 ‘16 :S9 “IR LUeD - SSOW LS
19 souor “N'gq'q ‘Wosuyos “HD Y
(WyYOY) DjIonuD vyapsoyoipuaddy

9661
“soy OAPI - YOOY UdsIOL 19 UasIa}od
“TW wniwpunjos puspjsosiup

6S/LS8I “E€TI-6IT *6 ‘PUOT 90S "A 90ld
- ‘ling sapiojpydaoua sndipo0j] (uy

BUTJODAWIOOSY

JINJVIIWT]

402

S19j}VM YSU

[B10 L -wg Jouul = pueyne jo
JO S}SVOD) 4SBOD SAMA
sarqyeo0’y SUOT}II][OD JO JIqUINN

isuny oY].

“SuOT}29{{09 YstuRg WWM [eep pajonb sanqesozy] oyL *(T “3ty) deur ayy pue (T 271483)

SIIPITEIOT JO ISI] 99S ‘SaIpPEIO'T *7X9} BY} Ul PoUOUTUIOD a1e X1J9}Se UL YIM poyIVUl saieds ‘uaseyuedo, ‘D pue qd Bleqgioy oq} Ul
paurejzuo0d aie pue si0yyne oy) Aq palpnys seq dARY SUOT}I9][09 [TY “S}SBOD YsTUBT WIJ Poo UO ISUNJ ULI JO SUOTIIAT[OD *Z 242L

403

a) 2) ‘y a EF ‘q

a2) =} “Ur ‘T ‘T ‘3 ‘q

I ‘3

rf a or ‘q

Lv ‘9p ‘bb ‘Ib

‘QE-VE “ZTE ‘TE ‘OE ‘PZ
Eee CC SI SI -FL-ez
Lv ‘OP ‘ph ‘Ib ‘Or
BUCO Ce TE. 8 Ce
Sie Clavie Ole 6c

8E “OE
eel ee Oea acc

br “Ip
OETPGr Ce 81-7

pr “SE “OE “Ss ‘7

br “6€ ‘ST ‘bl

tC EEC

Ge 0€ V1

gs

69

61

El

O€

a4

8V

cl

IT

16

cl

(a6

CLOT “E€S6I -OS “JOM fURD - WY “Md
(‘wyYyoy) véafignqni sisdosodsouay

br6l “60P :T BIMOTIE.F
- Japury pwyoy sisdosodso1ay

886I “LZE :ZE UOXEIODA| - [eUILZ Jo
souof “Dg vInpvs sisdosodso11a7) x

6961 “801 :08 “Yodey sappe.j - 1prumyog
‘| Sapiosodsoajd pyjasavydsoqivy

6961 ‘OI-6 :L
BINQUITASIPY ZINYISIMVN JEN - Ips
‘| Sapiouvydsojda] pyjasavydsoqivy

L861 “EL

88 ‘0G [OSAP] WIg‘sueI], - UeUISSOY

J9 sjonuresg ‘syst ‘¢°q ‘iMag

“TOW “A CulYyoy) summons viuosdn) x

C661
‘TSE-OPE (IL “WOM Lf uRD - souos ‘DN’ gg
19 YOOY UIsIEL DIVILUJUI DNULOIIOg

6861 “E8IT
‘LO “JOR fueD - souof “D' g°7A
19 YOY ussier Dosvskyd onyfig

wi ‘q Or ‘br ‘6E ‘S ‘7 61

yes bbe aa og eer ade | C37 6
Tere Oe Cc

rm ip Se Meee aes a 6k Clee ee eee Or

Ly ‘9b ‘ph ‘SE “PE
LE OCeSC. VORCE- SI

ny ‘u ‘ft “8 “q “ST “pl OL 8 “9-4 “@ 66
SE I

ja 13g OT ‘6 ‘8 ‘9 ‘S ‘7 LZ
ies Z I

tr I

404

Sc

6v

a

17

OS

9¢

997-S9T :97
‘IeJW10g - SSOW ‘L's 32 SOL “DN G'|A
‘YOY UsBleL DISaAIg DYJaIyUuaoIH

'

8961 “8S 18 “S9D
‘jog Yyosiq Jeg - ‘WYoY vp21un] DuCIT

6861 “S8IT :L9 “Og fuURD - souof “Hdd 39
yoy uasier supsiapu viavydsoMds

7ZOL “8HZ ‘8
woTyeq-ulplog snl ney 10g" [qZNON
- “ULIpIIM DUILDU D4Lodso]]040D

7861 “€OI-ZOT
-€Z “udg-d0g [OOAP] SURIT, - Fyeqny,
19 WISeyeN joan] v1éodsopjo410D

7961 “OTT “SL SPD Og YOsI 19g
- ‘wyyoy (Jepurq) 04290] psodsojjo10D

€961 ‘8OE-LOE :9 PISIMpoH BPAON
- ‘W]YOY vsojapyo visavydsojanyD

6861 ‘PIS ‘pe UoxRIOd\| - Leg “q'W
(ysrey “d) Djooiyddy vIspUojavY) x

ee ee ee ee ee ee SS SS ee

ponuryuos *Z 3fqey

405

n 4 UW of a ‘3 a

1°U I ‘8 ‘q

OI ‘8 “9 ‘¢ ‘Z

Ot vesht OF CH 11 -¢

Lv ‘vb ‘Ip “6€ ‘LE
COVE Ot LG Yee
Oc 81--9T P10 -<
Lp ‘9 ‘bb

“OV-TE ¥C-CC- GL SI
Seri TeOl pyc

br “9€ “bE

CE OS. bc. Ce GY]

6€

br ‘6E ‘IZ

Tr ‘6E

"SE “9E-PE “ZE ‘TE ‘67
CC VC PESOMS eT

LE

O€

8S

66

ch

87

5G

8V

(Ge

LL

(&6

Le

Ol

E861 ‘OIZ :L8 ‘90S uurT f10g - sso
‘LS 10 wosuyor “DY ‘sous “9° g'Y
(wYoy) pipjngn vjjauakaupyoy

C861 “E61 *16 “90g UUT] f10g - “MsymeH ‘Gq
(Jepur]) DuDW DYyaYyronnaisyosary

8561 “bh ‘PL ‘90S 19g [TOYO eYSTIA’f
- uosuyof “MA 'L (QQUD “Mf 19
Qquid) véasyasoipau sisdoianydsojopyy

pre! “TI *T BIMOTIE]
- lopury] Dyojnoipuaddp visapydsojvyy

C861 “OTP :pZ UOXRIODA|
- OD] nsuas “WTYOY-"UyOA 19 “WyTyoY
(4pruryog *[) vsoosia piaydivsoj0H x

E861 “O07 :08 ‘90 TOOK] Ig ‘sues ],
- oWOND 19 SSOP LS ‘souor “DH'G'q
(WyYOy) dsafizjna viaydivsojoyy

9861 “90P :€6 “90S “UUIT “f10g - sauos
‘O)'a'_ 30 ueuey “yD vulos vuasyoy

1961 ‘88-L8 ‘€ PISIMpaH PAON
- “W]YOY vioydosajvja Duasyoy

br ‘6€ ‘€7 6

a7
[ “q Ze 0S 66-82 S577 Or

pr ‘6€ ‘LE ‘SE ‘TE “OE
ySe ees “60. SCO. SI. Olec Os
ial od i I
Sis 9¢€ ‘SE ‘OE ‘BT El

tr ‘6€
EE-CE CCA GES ee 61

br “6€
13. 1-3 <q CECE STE LOL o> ¢ 87
q i I

406

Ol

8V

as

O1

1@

O€

6861 “O80€ :L9 JOR fURD
- apAH ‘Q' Jaaousunu diavydsouLuvfy x

v86I “68€
QZ UoxRIOdK] - SOUOLS “NH ' G7 19 YOY
UIBIGL VDLIDUAIDOUST] DIYUOMINT

9161 ‘6STZ 3 “90S JOA Wag sues]
- ‘Payng “yO DjooIonf viyLOMNT

9S6I ‘ITh °6€ “90S TOdA WIG
“SueI], - WOSTIM “WI 27D{ful DipurT

C661 “VIOT-IIOT -€L JOM LUeD - SoUOs
‘O'g'7 19 YOoy ussier (soUOS “DH ' gy
19 YoY uasier) DIIpuNsS D1OdsOIDUIT

Z96I ‘SOT ?Sp ‘90S TOOK Ag“ sues],
- squor “9 'q'q DI18Djad DiuavydsojdaT

prel “Ely :1 BIMopey
- Jopury] supwavio visavydsojdaT

C661 “6SST-OSST ‘ZL 0G [ UeD
- SSOW ‘LS 12 souog “O' a’ A ‘Wosnx “W
(wyYOY) visaaunos2 sisdosodsynvy]

Sg a ee ee

ponunuos ‘7 3[qe

407

i ea 8 eI as ‘q

n = ‘UW T op at ‘3 ‘q

11 ‘3 ‘q

LY

‘ph “6€ “9E “SE ‘ZE ‘OE
SEM GG EOE a Alen
br ‘Tr ‘Ib ‘SE

‘OE-VE ‘ZTE ‘OE ‘bZ ‘61
LEE Ch OL-8- Se

STE?

Shon as

vb

WW -Ce OE 61> SI

br “6€

pr “6€ “9€-rE
come. 81-9l a?

Ae

801

IT

4

6¢

61

BL

SY

€86I “907 :L8 ‘90g UUTT f10g
- SSO “L'S§ 39 UOsUYOL “Hy ‘souor
C ‘O'aA'A Cwypyoy) 018119 vsodsoiasan

E861 “907 :L8 ‘90g UU] [10g
- SSOW ‘LS 19 UOsUYoL “HY ‘souor
68 ‘Oa A CWyTYOy) vww09 vsodsoiasan

p96I °L6
‘Lp “00G [OSA] WAg‘sueI] - souor
c ‘aA Dinunuvjs19 viuapydsoinon

7961 “60h ‘hp PISIM
0 -P9H PAON - “WTYOY viowi0u2 s1oN

L861 “SLO *$9 “OG fURD - “WTYOY-"UNTOA
0 }9 “WYO avissyjvy) vuLvssvpyy.

6861
“€8IT :L9 “JOG fs uUeD - souos “N'gG'g
0 }2 YOOY uss1el DJVsuD stuDjOsUDP

9961 ‘SPS ‘IT PISIMpay BAON - “[RARD “YW
0 (‘WIYOY) Dwissisuo] viodsoutuDpy,.

9961 ‘8PS ‘IT PISIMpoH BAON - [RAPD “YW
0 (WYO) viv1Wdk{jv9 viodsouruvp

408

| ‘3 ‘q

Ot

ZE “PT

Op-8E “9E-PE “TE
POV cl 950
Lv ‘9b ‘bb “6E-VE
SOC SURI. SIP
EL. OLS SG ac
Lv ‘vb ‘Ov “9€ ‘ZE
Fi Gag) | BE he NES

6¢

6€

PEOOS «SE ce OE
Ge COSS i Sie VEC

Be

88

67

ov

91

tL

9¢

(63

CC

LA

vi

6861 “LIS ‘8 OGL IPION - Yooy
uasier DIpquvqiuids vaodsimay

O96T ‘TEE :Z PISIMPaH PAON - “WTYOY
(quYyOH) siuasuponb vsodsimay

C961 ‘6IE :9 PISIMpPOH
BAON - “WyYOyY vyvazjId vaodsimay

prel ‘Oly :T BIMopey
- Jopury vwyiuDwu diodsmuay

8961 “ZOE :ST “Qe T [Ol Je O19g
‘Iqng - Tyeqny], pyuazps vsodsmay x

I86I “LIE€T -6$ “O_ LueD
- ‘wyyoy vds19 piodsmay.

P86 ‘OCT LZ “Je 10g
- SSO ‘],'S 19 WOSUYOL “HY ‘souosr
‘O'a@'a (WTYOy) vonbso} vyaiuipuoC

E861 “PSE :9T “IRIN I0E
- SSOP “LS 30 UOsUYOL “HY ‘souos
‘OQ g'q Duassisuopjidpy vsodsvjsorO x

ponutjuos *7 3[qey

409

5 i | ey ‘Y ‘3 ‘q

OP ‘7h ‘Ih ‘6E “8E
‘9E-PE “ZE “OE “67
SG o- Viet 9 °C. 7

6€ “SE

Iv
bE{S6 SC Se. piael

Otc

Ors 6c ¢

VE 60 Ce Sled

vv

br “Th-8E
OEE 106 Sz
VGeLi stl ace veG

6v

0@

OI

(Z6

$6

(47

0¢

Ic

Iv

vs

L961 ‘T6¢ ‘d ‘JowRID ‘saremsy
pue sues00 UI IsuN, - soysny “DD
(Jopury) wno18pjad wniuodsoKjaig

C861 ‘OCh ‘PZ UOXRIODI]
- IPIUIYIS | vu1uvU Dsodsojnuny

O96I “LrE ‘Lr
‘Jog fUTY - BOOP ‘LY 10 sIdXI
(WYO) vjvydad049DU DIDUAaLAID

7961 “EIZI ‘Oh “JOM sf URD - nesIOW= “W
JO NRO] SNJWIINAD sadKuosajsy

BUNODAWIOIONOG

O96I “OEE :Z BISIMpoH PAON
- ‘WyYoy vsouidsiquv viodsopadioj

prel “Elp ‘1 BIMmopiey
- JOpur] SLDWaDs.O DuYNsavyds

6961 “Z9T :7S “90g
‘JOOAP[ Wig suey - uoIeT “yy ie
soot “9 gq'y Mjo21Iusy vyjakioavs

0961 ‘PEE :7 BISIMpoHx] PAON
- “WTYOY vjyv]]a1s vaodsnuay

410

‘Ly 9b
‘ph ‘Ip “6€ ‘BE “SE “CE
te! SeetCchl Vie Ol <c

6cac

vv

1“q yy 6c te OE

Ly

‘Ob ‘bh ‘TP “BE-CE “OE “67

fog oe [el ee CC Cl ale eo Pee

I rr ‘6E “SE ‘S ‘7

OV

8cl

ve

Il

86

O¢

C

C661 “7SE ‘eS

UOXEIOIAPY - “WYYOY- ULAJOA 1°
“‘wWyYyoy (a100W L'a 3 sIah3W)
wnjjauojjodojp wnippjI0YyI14 TL

IL61 “Phe
:€Q PIBO[OSA| - WOOW La 1?
slokay| wnsodspyov wunipojIOYyIUT

7861 “€OI-ZOT
€Z ‘udg90g JOOP] SURI - FYRQNL
19 INISeyeN Vj0IIN] paplowusis

prel ‘SOb-POr ‘1 BIMoted
- Jopury siiqvjoads sarkuigiO

C96I ‘“8ET :9P “90S
[OOK Wg suely - sof “D' ACaA
(uosuyor * ML) 09280]ad skjopouopy

prel ‘cOr :T PIMoptey
- Jopury siupwavso vIpo}diq

SO6T ‘TOI :1Z ‘Ja “[OoAU “90g “ISOUIN “TINE
- uandeny (eplod) sap10jn41o} wn110dsonjriq

poenutuos “7 s[qeL

411

| sr oF ‘q

ae

jelet

n ‘J af ‘3 ‘q

9€ “SE “CT ‘7

vr

sSECe CC VG EC Gee

Lr ‘9P

SL CleGe CC O61 El

GCTe. Ce
ce Cc moll. Lao av

Ip

él

VC

a

6P

I?

el

6€

OI

6S61 “PLS :1¢ eISO[OOKp
- SIBAZJI 19 DIOOW “LY VSssiugia DIN’

C96I “8EEP
‘PST “LOS peoy ssouveg “pqaH “YO
- Jons0q vuuuDUu vsodsyM3ig

BUNODATHOIpIseg

£961 “SEIT ‘Ip Jo fueD
- NOISEISLUY WNLIDA U01L3]DZ
€961 “9EIT

‘Ip JOG fueD - nolsejseuy
(1opUlT) wnwijLIDU U01La]DZ

S861 “OCh :pZ UOKRIOKW

- PITY “| wnjoujsuod wnipoyjdI0YoU T

412

Acknowledgement
The authors wish to thank Professor E.B. Gareth Jones, University of Portsmouth,
for reviewing the manuscript. The senior author express his gratitude to E.B. Gareth
Jones for his agelong interest and support in the study of marine lignicolous fungi
from the Danish coast. An interest and support without which this work would not
have been possible.

Literature

a. _ Ellis, M.B. (1971). Dematiaceous Hyphomycetes. CMI, Kew, 608p.

b. Farrant, C., Hyde, K.D. & Jones, E.B.G. (1985). Further studies on
lignicolous marine fungi from Danish sand dunes. Transactions of the British
Mycological Society. 85: 164-167.

c. Hermann, F. (1968). Danmarks Natur (havet) - Hydrografi. Politikens Forlag,
Kobenhavn. 3: 24-47.

d. Hohnk, W. (1955). Studien zur Brack- und Seewassermykologie V. Hohere Pilze
des submersen Holzes. Verdffentlichungen des Instituts fiir Meeresforschung in
Bremerhaven. 3: 199-227.

ec: Johnson, R.G., Jones, E.B.G. & Moss, S.T. (1991). Histochemical and electron
microscope studies of ascospore ontogeny in three marine fungi with
cleistothecia: Amylocarpus encephaloides Curry, Dryosphaera navigans Koch et
Jones and Eiona tunicata Kohlm. Botanica Marina. 34: 229-239.

f: Jones, E.B.G., Koch, J. & Moss, S.T. (1995). Limacospora gen.nov. and a re-
evaluation of Ceriosporopsis cambrensis. Canadian Journal of Botany. 73: 1011-
1014.

g. Koch, J. (1974). Marine fungi on driftwood from the west coast of Jutland,
Denmark. Friesia. 10: 209-250.

h. Koch, J. (1989). Remispora spinibarbata sp. nov., a marine lignicolous
ascomycete from Denmark. Nordic Journal of Botany. 8: 517-520.

i Koch, J. & Jones, E.B.G. (1983). Vedboende havsvampe fra danske kyster.
Svampe. 8: 49-65.

j. Koch, J. & Jones, E.B.G. (1984). Lulworthia lignoarenaria, a new marine
pyrenomycete from coastal sands. Mycotaxon. 20: 389-395.

k. Koch, J. & Jones, E.B.G. (1986). Ceriosporopsis sundica, a new lignicolous
marine ascomycete from Denmark. Nordic Journal of Botany. 6: 339-344.

ie Koch, J. & Jones, E.B.G. (1989). The identity of Crinigera maritima and three
new genera of marine cleistothecial ascomycetes. Canadian Journal of Botany.
67: 1183-1197.

m. Koch, J., Jones, E.B.G. & Moss, S.T. (1983). Groenhiella bivestia, gen. et sp.
nov., a lignicolous marine fungus from Denmark. Botanica Marina. 26: 265-270.

n. Kohlmeyer, J. (1968). Danische Meerespilze (Ascomycetes). Berichten der
Deutschen Botanische Gesellschaft. 81: 53-61.

o. Kohlmeyer, J. & Kohlmeyer, E. (1979). Marine Mycology. The higher fungi.
Academic Press, New York, 690p.

p. Kohlmeyer, J. & Volkmann-Kohlmeyer, B. (1987). Marine fungi from Aldabra,
the Galapagos, and other tropical islands. Canadian Journal of Botany. 65: 571-
582.

q. Kohlmeyer, J. & Volkmann-Kohlmeyer, B. (1991). Ilustrated key to the
filamentous higher marine fungi. Botanica Marina. 34: 1-61.

SS.

ZZ.

413

Lind, J. (1913). Danish fungi as represented in the herbarium of E. Rostrup.
Gyldendalske Boghandel-Nordisk Forlag, Copenhagen. 659 pp.

Petersen, K.R.L. & Koch, J. (1996). Anisostagma rotundatum gen. et sp. nov.,
a lignicolous marine ascomycete from Svanemgllen Harbour in Denmark.
Mycological Research.

Read, S.J., Jones, E.B.G., Moss, S.T. & Hyde, K.D. (1995). Ultrastructure of
asci and ascospores of two mangrove fungi: Swampomyces armeniacus and
Marinosphaera mangrovei. Mycological Research. 99: 1465-1471.

Rees, G., Johnson, R.G. & Jones, E.B.G. (1979). Lignicolous marine fungi from
Danish sand dunes. Transactions of the British Mycological Society. 72: 99-106.
Rees, G. & Jones, E.B.G. (1985). The fungi of a coastal sand dune system.
Botanica Marina. 28: 213-220.

Rostrup, E. (1884). Underjordiske svampe i Danmark. Meddelelser fra den
botaniske Forening. 1: 102-106.

Rostrup, E. (1888). Mykologiske Meddelelser. Meddelelser fra den botaniske
Forening. 2: 84-93.

Schaumann, K. (1975). Okologische Untersuchungen tiber héhere Pilze im Meer
-und Brackwasser der Deutschen Bucht unter besondere Berticksichtigung der
holzbesiedelen Arten. Verdffentlichungen des Instituts fiir Meeresforschung in
Bremerhaven. 15: 79-182.

Schmidt, I. (1974). Untersuchungen iiber héhere Meerespilze an der Ostseekiiste
der DDR. - Natur und Naturschutz in Mecklenburg. 12: 1-148.

Yusoff, M., Jones, E.B.G. & Moss, S.T. (1995). A reappraisal of the taxonomy
of the genus Ceriosporopsis Linder based on their ultrastructure. Canandian
Journal of Botany. 72: 1550-1559.

Yosoff, M., Koch, J., Jones, E.B.G. & Moss, S.T. (1993). Ultrastructural
observations on a marine lignicolous ascomycete: Bovicornua intricata gen. et sp.
nov. Canadian Journal of Botany. 71: 346-352.

SEM micrograph. Fig. 2. Marinosphaera mangrovei. Ascospore covered by a layer of
fibrillar material (after 15 min settlement on the membrane). Scale bar = 5 pm.

414

Light micrographs. Figs 3, 4. Capronia ciliomaris. Fig. 3. Ascoma with a
membraneous peridium. Fig. 4. Ascospore surrounded by radiating appendages forming
knots at the tips. Figs 5-7. Marinosphaera mangrovei. Ascospores. Fig. 8. Massarina
thallasiae. Ascospore enclosed in a mucilaginous sheath. Fig. 9. Ondiniella torquata.
Ascospores with polar rigid appendages and a tubular annulus around the septum. Fig.
10. Nautosphaeria cristaminuta. Ascospores with polar and equatorial appendages. Fig.
11. Lautisporopsis circumvestita. Ascospore covered by an irregular mucilaginous
sheath, protuding at the poles and around the septum. Scale bars: Fig. 3 = 50 um, figs
4-11 = 10 pm.

MYCOTAXON
Volume LX, pp. 415-428 October-December 1996

LICHENICOLOUS FUNGI FROM THE
WESTERN PYRENEES, FRANCE AND SPAIN.
II. MORE DEUTEROMYCETES

Javier ETAYO

Navarro Villoslada, 16, 3° dcha
E-31003 Pamplona, Spain

Paul DIEDERICH
Musée national d’histoire naturelle

Marché-aux-Poissons
L-2345 Luxembourg, Luxembourg

Abstract: 12 species of lichenicolous deuteromycetes are recorded from
the western Pyrenees. 4 species are described as new: Hainesia
pertusariae Etayo & Diederich (on Pertusaria sp.), Macrophomina
pseudeverniae Etayo & Diederich (on Pseudevernia furfuracea),
Milospium lacoizquetae Etayo & Diederich (on Cladonia parasitica) and
Sclerococcum parmeliae Etayo & Diederich (on Parmelia s.1.). The
genus Marchandiomyces is recognized to include at least two species: M.
corallinus (Roberge) Diederich & D. Hawksw. with pinkish, initially
immersed conidiomata, and M. aurantiacus (Lasch) Diederich & Etayo
comb. nov., with orange, more or less superficial conidiomata.

Introduction

This paper represents the second of our series dealing with lichenicolous
fungi from the western Pyrenees. Like the first part (Etayo & Diederich
1995), we study here a number of lichenicolous deuteromycetes.

Material and Methods

Most specimens studied are located in the private herbaria of the authors, type
specimens in K, M, MA-Lichen and UPS. Microscopical examination of
herbarium material has been done at a magnification of x1000, measurements
in water, and illustrations in water or Congo-red.

416

Results and Discussion

Ampullifera foliicola Deight.

France: Pyrénées-Atlantiques, S of Tardets-Sorholus, Gorges de Kakouetta, on
Byssoloma subdiscordans (Nyl.) P. James, 26 July 1990, P. Diederich 9462; ibid., on
B. leucoblepharum (Nyl.) Vainio, 31 Jan. 1993, J. Etayo 12169. - Spain: Catalunya,
Girona, Oix, Riera d'Oix, 500 m, on B. subdiscordans, Febr. 1991, J. Etayo 5726;
Guiptizcoa, Jaizkibel, on B. leucoblepharum, 15 Nov. 1991, J. Etayo 6249; Navarra,
E of Lumbier, Foz de, Arbaytn, on B. subdiscordans, 21 July 1991, J. Etayo 5120;
ibid., 7 May 1987, J. Etayo s. n.

Hawksworth (1979) reported this species from Brazil and Ghana on folii-
colous lichens. The exsiccata Santesson Fungi Lich. Exs. 51 represents
material from Madeira on Byssoloma subdiscordans. Sérusiaux (in press)
mentions its presence on Byssoloma subdiscordans in Madeira and in the
French Pyrenees. The fungus seems to be not rare in regions where
Byssoloma species exist.

Bachmanniomyces uncialicola (Zopf) D. Hawksw.

Spain: Navarra, Urroz de Santesteban, pantano de Leurtza, 900 m, on Cladonia
furcata (Huds.) Schrader, 13 Febr. 1994, J. Etayo 12173.

Hawksworth (1981) recognized this species as a parasymbiont on the
Cladonia uncialis (L.) Weber group, but mentioned a few doubtful literature
records on other species of Cladonia. This is the first record on Cladonia
furcata and also the first record of the species from Spain.

Fusarium peltigerae Westend. & Wallys

Spain: Navarra, Baztén Valley, between Errazu and Izpegui, on Parmeliella testacea
P. M. Jgrg., 440 m, 6 March 1994, J. Etayo 12227.

Our specimen fits well the description of this species given by
Hawksworth (1979). It forms pinkish to orange sporodochia, slimy when wet
and irregular when dry. Conidiogenous cells are 17-27 x 2.5-3 um, conidia
curved or straight, hyaline, 1-3 septate, 85-117 x 3-4 um. Dr Dagmar Triebel
kindly drew our attention to the correct author citation of the species name
(given as F. peltigerae Westend. by Hawksworth 1979).

The species was known from the type collection from Belgium and from
Sweden (Santesson 1993), both on specimens of Peltigera rufescens (Weis)
Humb. Our collection from the Pyrenees was growing as a saprophyte on an
unhealthy thallus of Parmeliella testacea.

417

This species was the only lichenicolous species of the genus recognized
by Hawksworth (1979), but we have seen at least three other unidentified
species of Fusarium with much smaller conidia growing on the thallus of
Pertusaria, Phlyctis and Parmelia in the Pyrenees.

Hainesia pertusariae Etayo & Diederich sp. nov. (Fig. 1)

Conidiomata lichenicola, cupulata, discoidea, semi-immersa, atrobrunnea, 80-150 um,
basi stromatica, pariete brunneo, cellulis elongatis. Conidiophora elongata, filiformia,
septata, hyalina. Cellulae conidiogenae enteroblasticae, integratae, sub conidiophorum
septis crescentes atque ramos laterales brevissimos formantes, hyalinae, leves.
Conidia hyalina, 0(-1)-septata, levia, filiformia, 14-22 x 1-1.5 um.

Type: Spain, Navarra, Urroz de Santesteban, pantano de Leurtza, in a Fagus
wood, on Pertusaria sp., 900 m, 13 Febr. 1994, J. Etayo 12699 (MA-Lich holotypus;
herb. Diederich, herb. Etayo isotypi).

Fig. 1. Hainesia pertusariae (holotype). A, cross section through a
conidioma. B, wall structure. C, conidiogenous cells and conidia. Scale bar =
10 um.

418

Conidiomata cupulate, discoid to slightly elongate, concave, semi-immersed
in the thallus of Pertusaria, dark brown, with a continuous or lobulate border,
80-150 um in diam; wall brownish, 15-20 wm thick, hyphal, composed of elon-
gate cells. Conidiophores arising from a basal stroma, 15-25 wm thick, com-
posed of more or less polygonal, hyaline cells. Conidiophores elongate, filiform,
simple or branched, septate, hyaline. Conidiogenous cells enteroblastic,
integrated, appearing as short lateral branches just below each septum of the
conidiophores, hyaline, smooth walled. Conidia hyaline, simple or indistinctly
1-septate, smooth-walled, filiform, 14-22 x 1-1.5 um.

Host: Pertusaria sp. (thallus), on shaded Fagus wood. The holotype
specimen is also infected by other lichenicolous fungi (Cornutispora
lichenicola D. Hawksw. & B. Sutton and a brownish myxomycete).

Distribution: Known only from the type locality in Spain (Navarra).

Observations: This is the first lichenicolous species of the genus Hainesia
Ell. & Sacc. The type species, H. lythri (Desm.) Hohn., has very similar
conidiomata and a similar conidiogenesis, but is distinguished by much shorter
conidia which measure 5-7.5 x 1.5-2 tum, and by a different ecology, the species
growing mainly on leaves (Sutton 1980).

Lichenoconium cargillianum (Lindsay) D. Hawksw.

France: Pyrénées-Atlantiques, SW of Larrau, near the road to Spain, on Usnea florida
(L.) Wigg., 29 July 1990, P. Diederich 9168; Pyrénées-Atlantiques, between col de
Suscousse and Arette ski station, on Platismatia glauca (L.) Culb. & C. Culb., 23
June 1992, J. Etayo 12677.

This species has originally been described on ‘Parmelia perforata’ from
New Zealand; the host has recently been identified as Rimelia cetrata (Ach.)
Hale & A. Fletcher (Triebel & al. 1995: 82). Hawksworth (1977) considers
Coniothyrium ramalinae Vouaux, described on Ramalina yemensis (Ach.)
Nyl. from Mexico, as a synonym of this species. Later Hawksworth (1981)
reports the species from the British Isles on the apothecia of Usnea florida.
Here we report it on Usnea florida and on Platismatia glauca. More speci-
mens on all these hosts are needed to understand if they are really conspecific,
or if several similar species are involved.

Lichenopuccinia poeltii D. Hawksw. & Hafellner

France: Pyrénées-Atlantiques, S of S'-Jean-Pied-de-Port, forét d'Iraty, 0.5 km S of
chalet Pedro, 1000 m, on Parmelia saxatilis, 27 July 1990, P. Diederich 9236; ibid.,
19 July 1991, P. Diederich 9685 & J. Etayo 5936, ibid., on P. sulcata Taylor, P.
Diederich 9686; Pyrénées-Atlantiques, Bois d’Astaquieta, between Occabe and
Sterencuby, 1200 m, on P. saxatilis, 18 April 1995, J. Etayo 12893; Vosges, Feigne
d’Artimont, 1.2 km NNW of Kastelleg, 1100 m, on P. sulcata, 14 June 1975, W. O.

419

van der Knaap & H. F. van Dobben s. n. (herb. Aptroot, herb. Diederich). - Spain:
Andalucia, Malaga, Estepona, Sierra Bermeja, subida a los Rales, 1235 m, on P.
sulcata, 18 April 1994, J. Etayo 12285, M. Casares, A. Gémez-Bolea & N. Hladun,
Navarra, W Pamplona, Sierra de Urbasa, P'® de Urbasa, on P. saxatilis and P. sulcata,
20 July 1991, P. Diederich 9640 & J. Etayo 5940; Navarra, N of Ochagavia, Iraty
wood, NNE of the lake, Nature Reserve "Lizardoya", on P. saxatilis, 19 July 1991, P.
Diederich 9668; Navarra, Espinal-Auzperri, on P. sulcata, 940 m, 4 Dec. 1994, J.
Etayo 12700. - Canada: British Columbia, Kispiox area, ca. 9 km NNW of Town, N
of Date Creek, 55°25’ N, 127°48’ W, 510 m, on Parmelia hygrophila Goward &
Ahti, 12 July 1992, T. Goward 92-302 & H. Knight (UBC, herb. Diederich).

This hyphomycete was known from Austria, Greenland and Scotland,
always growing on species of Parmelia s. s. (Hawksworth 1990). We report it
here as new for France, Spain and Canada. It seems to be widespread, and
rather common in areas with an oceanic climate.

Macrophomina pseudeverniae Etayo & Diederich sp. nov. (Fig. 2)

A Macrophomina phaseolina cellulis conidiogenis latis et phialidibus angustis minus
distinctis differt.

Type: Spain, Huesca, road Sabifidnigo to Boltafia, Puerto del Serrablo, Sierra del
Gabardon, km 39 near Laguarta, 1200 m, on Pseudevernia furfuracea, 5 July 1993, J.
Etayo 12597 (MA-Lich holotypus; herb. Etayo isotypus).

Mycelium immersed, hyaline. Conidiomata pycnidial, separate, globose,
dark brown to black, immersed, unilocular, 130-170 um in diam.; wall com-
posed of a brown textura angularis, paler inside, darker near the ostiole; cells
thin-walled, 6-18 x 5-15 um. Ostiole central, circular, slightly papillate.
Conidiophores absent. Conidiogenous cells enteroblastic, discrete, determinate,
ampulliform, hyaline, smooth, often indistinct, 7-10 x 6-7 um. Conidia hyaline,
aseptate, obtuse at each end, sometimes slightly constricted at the point of
attachment, straight, cylindrical, ellipsoid or sometimes irregular in form, thin-
walled, smooth, with small guttulae, (14-)16-22(-26) x 6-9 um.

Host: The species is parasitic on the thallus of Pseudevernia furfuracea
(L.) Zopf growing on Pinus sylvestris. Infected parts of the thallus get pinkish
brown.

Distribution: Only known in more or less continental areas of northern
Spain (Huesca and Navarra) where the species seems to be not rare.

Observations: The phialides in the new species are narrow and some-
times indistinct. In the type species of the genus, Macrophomina phaseolina
(Tassi) Goid., the phialides are very distinct, with a wide aperture, and the
conidiogenous cells are narrower (5-13 x 4-6 um); that species is plurivorous
(Sutton 1980).

Additional specimens (all on Pseudevernia furfuracea): Spain: Huesca, Aragiies
del Puerto, 26 Aug. 1991, J. Etayo 770; Huesca, Piedrafita, track to Pefia Telera,

420

1400 m, 4 Aug. 1993, J. Etayo 12598; Navarra, Sierra de Ill6n, monte Belbin,
Navascués, 2 km from Castillonuevo, 1250 m, 11 June 1994, J. Etayo 12599, 12600
(herb. Etayo, herb. Diederich); Navarra, Belabarce, 25 July 1987, J. Etayo 3209.

. C

Fig. 2. Macrophomina pseudeverniae (holotype). A, immersed conidioma
showing textura angularis of the wall. B, conidiogenous cells with conidia. C,
conidia. Scale bars: A = 50 um; B, C = 10 pm.

Marchandiomyces corallinus (Roberge) Diederich & D. Hawksw. s. I.

The genus Marchandiomyces Diederich & D. Hawksw. has been created to
include the fungus previously known as Illosporium corallinum Roberge. The
conidiogenesis, and more generally the nature of the different cells visible
inside one fructification, are very difficult to interpret (Hawksworth 1979:
236, Diederich 1990). Dr Rosalind Lowen informed us about her most

421

interesting discovery that dolipores have been found in American material of
M. corallinus (s. 1.!), showing that it represents a basidiomycetous anamorph.

Hawksworth (1979: 237) correctly noted that the species (i.e. M.
corallinus s.1.) presents some variation in the colour of the fructifications,
and that the orange and the pink specimens could well belong to two different
species. Field studies in Luxembourg and in the western Pyrenees allow us to
affirm that at least two different species are involved:

1. Marchandiomyces aurantiacus (Lasch) Diederich & Etayo comb. nov.

Basionym: //losporium aurantiacum Lasch, in Rabenhorst, Fungi Eur., cent. I, n°
74, 1859.

Type: Germany, in lichenibus trunci Pyri mali pr. Driesen, G. W. Lasch, Rabenh.
Fungi Eur., cent. Ino. 74 (K, M, UPS - isotypes!).

Exsiccatae: Fuckel Enumeratio Fungorum Nassoviae, ser. | no. 206 (M! sub J.
aurantiacum). - Fuckel Fungi rhenani, no. 239 (M!, sub I. aurantiacum), 241 (M!,
UPS!, sub J. roseum Fr.). - Rabenhorst Fungi Eur., cent. I no. 74 (K!, M!, UPS!, sub
I. aurantiacum).

A light orange species (Kornerup & Wanscher 1984: 5A4, the coiour
corresponds to recent herbarium specimens) with fructifications always
superficial, growing on macrolichens in the Xanthorion communities, mainly
on Phaeophyscia, Physcia and Xanthoria; this species is quite common in the
industrialized and polluted south-western part of Luxembourg, but has not
been found in the western Pyrenees. It is often found together with Hobsonia
christiansenii. We studied three isotypes of Illosporium aurantiacum Lasch
(K!, M!, UPS!): the specimens are now in a poor condition, and only a few
superficial or slightly immersed fructifications could be seen over a thallus of
Physcia adscendens. In the original description the colour of the fungus is
said to be orange (‘aurantiacum’); due to the age of the specimens, the
fructifications are now pale, which is typical for old specimens of this species.
We conclude therefore that the type specimen of /. aurantiacum belongs to
the species with orange fructifications, which has therefore to be called
Marchandiomyces aurantiacus.

Hosts: We have seen material on Phaeophyscia orbicularis (Neck.)
Moberg, Physcia adscendens (Fr.) H. Olivier, P. caesia (Hoffm.) Firnr., P.
semipinnata (J. F. Gmelin) Moberg, P. stellaris (L.) Nyl., P. tenella (Scop.)
DC., Xanthoria candelaria (L.) Th. Fr. and X. parietina (L.) Th. Fr.

Distribution: We have seen specimens from Austria, France, Germany,
Luxembourg, Spain and Sweden.

Additional specimens (Luxembourg specimens of this species published by
Diederich 1990, sub M. corallinus, are not repeated here): France: Gard, near Pont du
Gard, on Physcia tenella, 13 April 1995, P. Diederich 12323. - Germany:
Wiirttemberg, Schwabische Alb, Winterlingen, Harthausen, 720 m, on P. tenella, 9
Nov. 1993, M. Heklau s. n. (STU, herb. Diederich). - Luxembourg: Kockelscheuer, on

422

P. tenella, 7 Oct. 1993, L. Reichling s.n. (herb. Diederich). - Spain: Andalucia,
Malaga, Ronda, pinsapar de la Cafiada del Cuerno, 1500 m, on P. semipinnata, J.
Etayo 12217, 12679; Navarra, Larra, Eskilzarra, near refugio, 1400 m, on P. caesia,
29 Aug. 1995, J. Etayo 12858; Navarra, alto de Erro, 800 m, on Xanthoria parietina,
18 April 1995, J. Etayo 12878. - Sweden: Sédermanland, Over-Jarna s:n, Bankesta
Gard., on P. tenella, 1 Jan. 1947, R. Santesson s. n. (UPS).

2. Marchandiomyces corallinus (Roberge) Diederich & D. Hawksw.

Type: France, on Physcia tenella, J. B. H. J. Desmaziéres, Pl. crypt.-Fr., Ed
fasc. 32 no. 1551 (K - isotype!).

Exsiccatae: Desmaziéres Pl. crypt. Fr., Ed. 1, fasc. 32 no. 1551 (K!). The speci-
men P. Diederich 12325 will be distributed in Santesson Fungi Lich. Exs.

A pinkish or reddish species (Kornerup & Wanscher 1984: 8A5, pastel
red) with young fructifications developing inside the lichen thallus, erumpent
and later superficial, often leaving holes in the lichen thallus, sometimes
spreading afterwards superficially over the lichen thallus, growing in less
polluted and more humid areas, generally in forests, often on Parmelia s. 1.
species; this species has been collected several times in the western Pyrenees
by the authors, but is not known from Luxembourg. Its colour is similar to
that of Hobsonia christiansenii Brady & D. Hawksw., but both species have a
different ecology and never grow together. The type specimen of J/losporium
corallinum Roberge (K!) belongs to this species, which has therefore to be
called Marchandiomyces corallinus. In the type specimen, numerous pinkish
fructifications develop inside the thallus of Physcia adscendens and later
become superficial; neighboring thalli of Xanthoria parietina are also
attacked. The original description (Desmaziéres, Pl. crypt. Fr., Ed. 1, fasc. 32
no. 1551, 1847) says that ‘Cet [/losporium nait dans la substance du support,
et la déchire pour devenir superficiel. ... Ils sont d’un rose carné, et recouverts
d’une trés-légére poussiére blanche.’, which corresponds perfectly to the
pinkish species collected in the Pyrenees.

Hosts: The species is known from Acarospora fuscata (Schrad.) Th. Fr.,
Flavoparmelia caperata (L.) Hale, Hypotrachyna revoluta (Florke) Hale,
Melanelia exasperata (De Not.) Essl., M. glabratula (Lamy) Essl., M.
panniformis (Nyl.) Essl., Melanelia sp., Parmelia omphalodes (L.) Ach., P.
saxatilis (L.) Ach., P. sulcata Taylor, Parmelina quercina (Willd.) Hale,
Parmeliopsis ambigua (Wulfen) Nyl., Parmotrema crinitum (Ach.) M.
Choisy, Pertusaria multipuncta (Turner) Nyl., P. pertusa (Weigel) Tuck., P.
pustulata (Ach.) Duby, Physcia adscendens, P. aipolia (Ehrh. ex Humb.)
Fiirnr., Ramalina fastigiata (Pers.) Ach., Usnea sp. and Xanthoria parietina.

Distribution: We have seen specimens collected in Austria, the Canary
Islands (Gomera), France, Norway, Spain and Switzerland.

Additional specimens: Austria: Oberésterreich, Donautal, Freizell, am Grat
gegeniiber Schloss Marsbach, on Acarospora fuscata, 1 Aug. 1993, F. Berger 6763

423

(herb. Berger). - Canary Islands: Gomera, Hermigua, mirador de El Bailadero, alt. 1010
m, on Usnea, 4 Aug 1994, J. Etayo s.n. - France: Finistére, S' Herbot, chateau du
Rusquec, on Pertusaria pertusa, 19 July 1954, R. Santesson 10256 (UPS); Pyrénées-
Atlantiques, SSW of Mauléon-Licharre, forét des Arbailles, S of S'-Just-Ibarre, vallon
du Bidouze, 0.5 km S of Arla, on Parmotrema crinitum, 27 July 1990, P. Diederich
9297; Pyrénées-Atlantiques, Orthez, route de Pau, on Flavoparmelia caperata, S.
Bahler s.n. (herb. Aptroot); Pyrénées-Atlantiques, vallée d’Aspe, near Urdos, on
Parmelina quercina, 1995, J. Etayo 12717; Pyrénées-Atlantiques, vallée d’ Aspe, near
Estaut, gorges de Pont d’Enfer, on Ramalina fastigiata, 7 Dec. 1993, J. Etayo 12676;
Pyrénées-Atlantiques, col d’Ibardin, La Redoute des Emigrés, on P. crinitum, 14
March 1995, J. Etayo 12702 & B. Marbach; Vaucluse, 2 km S of Gordes, col de
Gordes, on Physcia aipolia, 21 April 1995, P. Diederich 12325, G. Clauzade, P.
Navarro & C. Roux. - Norway: Oppland, Vaga, Veomoan, on Melanelia panniformis,
7 Aug. 1985, R. Santesson 31265 & P. M. Jorgensen (UPS); Sor-Trondelag, Brekken
herred, near Botnlia, 4-5 km SSW of Brekken, on M. exasperata, 31 July 1986, R.
Santesson 31546c (UPS). - Spain: Catalunya, Girona, on Melanelia sp., Oct. 1989, B.
Matesanz s.n. (BCC, herb. Diederich); Navarra, Narbarte, on P. omphalodes, P.
saxatilis and P. sulcata, 3 Sept. 1991, J. Etayo 13107, 13108, 13109; Navarra,
Roncesvalles, Lindux, on P. sulcata, 15 Sept. 1991, J. Etayo 6013; Navarra, Oronoz-
Mugaire, Sefiorio de Bértiz, on Hypotrachyna revoluta, 13 Nov. 1992, J. Etayo 3734;
Navarra, 20 km E of San Sebastian, Col d’Ibardin, on F. caperata and M. glabratula,
22 July 1991, P. Diederich 9758 & J. Etayo 5980; Navarra, valle de la Ulzama,
Alcoz, on Pertusaria pustulata, May 1993, J. Etayo 12703; Navarra, between
Roncesvalles and Valcarlos, km 55.5, on P. multipuncta, 19°5, J. Etayo 12883. -
Switzerland: Wallis, Bourg-S'-Pierre, nahe dem S'-Bernhard-PaB, Waldweg an der
Ostflanke des Créta de Vella, im Foret du Milieu, on Parmeliopsis ambigua, 10 June
1991, C. Printzen 2138 (M).

3. Marchandiomyces sp.

A number of specimens cannot be included in a convincing way in one of
these two species; they are either too old or in a poor condition, or they might
belong to different undescribed species. Two specimens with brownish
(cinnamon) fructifications (Kornerup & Wanscher 1984: 6D6) have been
collected in Luxembourg (Diederich 8442) and in Sweden (R. Santesson
48.502d, UPS); they represent most probably a third species, but we prefer to
await the discovery of additional specimens before describing it as new.
Several North American specimens on Cladonia stellaris (Opiz) Pouzar &
Vezda, Lasallia papulosa (Ach.) Llano and Rhizoplaca chrysoleuca (Sm.)
Zopf (Harris 16637, 16638, 16642 NY) have pale yellowish or brownish
yellowish fructifications (Kornerup & Wanscher 1984: 4A4); they don’t seem
to belong to one of the two named species; it is important to note that the
specimen on Rhizoplaca (Harris 16642) has been used by Dr Lowen to study
the dolipores, and that her discoveries are thus not based on M. aurantiacus
or M. corallinus, but on an apparently undescribed species of
Marchandiomyces. !

424

Milospium lacoizquetae Etayo & Diederich sp. nov. (Fig. 3)

Fungus lichenicola. Conidiophora in sporodochiis densis convexis atrobrunneis,
rotundatis, (30-)50-100(-120) jm, semi-macronemata, simplicia, determinata.
Cellulae conidiogenae monoblasticae, integratae, terminales, determinatae. Conidia
variabilia, verruculosa, brunnea, 7-14(-16) x 6-10(-12) tm, cum maculis
obscurioribus.

Type: Spain, Navarra, Oronoz-Mugaire, Sefiorio de Bértiz, 360 m, 30TXN1579,
in a Fagus wood, on Cladonia parasitica, 4 Jan. 1994, J. Etayo (MA-Lich holotypus;
herb. Etayo isotypus).

Colonies forming convex, blackish, rounded sporodochia, (30-)50-100
(-120) um in diam.; mycelium immersed in the host thallus. Conidiophores
semi-macronematous, agglomerated in very dense sporodochia. Conidiogenous

\/ :

Fig. 3. A, Milospium lacoizquetae (holotype), conidia. B, Sclerococcum
serusiauxii (J. Etayo 1458), conidia. Scale bar = 10 um.

425

cells monoblastic, integrated, terminal, determinate. Conidia variable in form,
brown, more or less lobed, apparently non septate, 7-14(-16) x 6-10(-12) um,
lobes (2-)3-5(-7) um, wall verruculose, unevenly thick, the thicker parts darker
than the rest of the conidia.

Etymology: The new species is named in honour of the neglected and
keen lichenologist P. Lacoizqueta (1831-1889), who lived in the valley of the
type locality.

Host: Cladonia parasitica (Hoffm.) Hoffm., mainly on the tip of
squamules. The fungus is parasitic, causing small brownish patches in the
host thallus.

Distribution: Known only from very shaded and well preserved valleys in
the north of Navarra (Spain).

Observations: This is the second species of the genus Milospium, the
other being M. graphideorum (Nyl.) D. Hawksw. (Hawksworth 1979). M.
lacoizquetae differs in the very small conidiomata, in the conidia which are
more lobed and in the different host (M. graphideorum grows on crustose
lichens with Trentepohlia as the photobiont) on which it forms brown sporo-
dochia. M. lacoizquetae also resembles Sclerococcum serusiauxii Boqueras
& Diederich in conidial shape and size (Boqueras & Diederich 1993), but the
conidial wall in the new species is much thinner and broken with many fis-
sures, the wall thickenings are not so evident and the conidia are not septate
(see Fig. 3).

Additional specimen: Spain: Navarra, between Errazu and Izpegui, on Cladonia
parasitica, 440 m, 6 March 1994, J. Etayo 12452 (herb. Etayo, herb. Diederich).

Pyrenochaeta xanthoriae Diederich

Spain: Navarra, W Pamplona, Sierra de Urbasa, P® de Urbasa, 20 July 1991, P.
Diederich 9630 & J. Etayo 5692; Navarra, Cilveti, 4 Dec. 1994, J. Etayo 12552;
Navarra, Fustifiana, Las Bardenas, caidas de La Negra, 600 m, 26 Febr. 1995, J.
Etayo s.n.; Navarra, alto de Erro, 800 m, 18 April 1995, J. Etayo 12880; Soria,
Calatafiazor, pista a La Fuentona, 1000 m, 27 Aug. 1995, J. Etayo s. n.

This species was previously known from Luxembourg (Diederich 1990)
and from Spain (Zaragoza) (Etayo & Blasco Zumeta 1992). All known
records are on epiphytic thalli of Xanthoria parietina.

Sclerococcum parmeliae Etayo & Diederich sp. nov. (Fig. 4)

Conidiomata lichenicola, sporodochia, rotundata, convexa, superficialia, griseofusca,
50-120 tm. Conidiophora semi-macronemata. Cellulae conidiogenae monoblasticae,
integratae, terminales, ellipsoideae, griseae vel fuscae. Conidia catenata, sicca,

426

acrogena, (0-)1(-2-)septata, ellipsoidea, grisea vel fusca, pariete rugosa, 7-9 x 3.5-5
um.

Type: Spain, Navarra, valle de Basaburia Mayor, between Aizdroz and Arrar4s,
track to Bergafie, on Fagus sylvatica, on Parmelia saxatilis, 550 m, 20 Sept. 1994, J.
Etayo 12688 (MA-Lich holotypus; herb. Etayo isotypus) (also present on P. sulcata
in the type collection).

Colonies forming discrete patches or forming a mass on the thallus of the
host; mycelium immersed, brownish. Conidiophores semi-macronematous,
aggregated into tufted, more or less rounded, convex, greyish brown, simple
to aggregated, superficial sporodochia of 50-120 um in diam., grey or pale
brown. Conidiogenous cells monoblastic, integrated, terminal, ellipsoid, grey
or brown, not clearly defined and the terminal cells acting in turn as conidio-
genous cells. Conidia adhering in chains, dry, acrogenous, (0-)1(-2)-septate,
ellipsoid, constricted at the septum or not, sometimes with one or two pointed
ends, grey to brown, ornamented, 7-9 x 3.5-5 um.

Hosts: On corticolous thalli of Parmelia s.1. (Flavoparmelia caperata,
Hypotrachyna revoluta, Parmelia saxatilis, P. sulcata, Punctelia subrudecta
(Nyl.) Krog and Rimelia reticulata (Taylor) Hale & A. Fletcher). As the hosts
are often in a bad condition and support other lichenicolous fungi, it is not
clear wheter S. parmeliae is saprophytic or parasitic.

Distribution: French and Spanish Pyrenees, common but apparently
overlooked.

Fig. 4. Sclerococcum parmeliae (holotype). A, chains of conidiogenous cells
and conidia. B, attachment of conidia. C, conidia (wall ornamentation repre-
sented on the left conidium). Scale bar = 5 um.

427

Observations: The new species is well characterized by the ellipsoid,
normally 1-septate conidia attached diagonally in columns. All other known
species of the genus have conidia which are either non-septate or more than
|-septate. The conidiomata are sometimes aggregated, forming brownish grey
masses on the thallus of the host.

Additional specimens: France: Pyrénées-Atlantiques, col de Lizuniaga, c. 5 km
from Bera de Bidasoa, on Flavoparmelia caperata, 5 March 1994, J. Etayo 12674 (herb.
Etayo, herb. Diederich); Pyrénées-Atlantiques, S'-Pée, 3 km from the village to Ustaritz,
150 m, on Hypotrachyna revoluta, 6 March 1994, J. Etayo 12682; ibid., on Punctelia
subrudecta, J. Etayo 12670, 12918; Pyrénées-Atlantiques, 5 km from Bera de Bidasoa
(close to the Navarra border), bois de Sare, col de Lizuniaga, 150 m, on Rimelia
reticulata, 14 March 1995, J. Etayo 12697; ibid., on F. caperata, J. Etayo 12698. -
Spain: Navarra, Oronoz-Mugaire, sefiorfo de Bértiz, 400 m, on F. caperata, 4 Jan.
1994, J. Etayo 12680, 12675; Navarra, Urroz de Santesteban, pantano de Leurtza,
900 m, on H. revoluta, 13 Febr. 1994, J. Etayo 12673; Navarra, valle de la Ulzama,
amongst Oroquieta and Saldias, km 7, 950 m, on H. revoluta, Febr. 1994, J. Etayo
12672; Navarra, valle de Basaburtia Mayor, amongst Aizdroz and Arrards, track way
to Bergafie, 550 m, on F. caperata, 20 Sept. 1994, J. Etayo 12671.

Sclerococcum simplex D. Hawksw.

France: Pyrénées-Atlantiques, S of S'-Jean-Pied-de-Port, forét d'Iraty, 0.5 km S of
chalet Pedro, alt. 1000 m, on Pertusaria pertusa, 27 July 1990, P. Diederich 9222:
Pyrénées-Atlantiques, S of Pau, W of Bielle, 0.5 km E of col de Marie-Blanque, on P.
cf. ophthalmiza (Nyl.) Nyl., 28 July 1990, P. Diederich 9329; Pyrénées-Atlantiques,
col de Lizuniaga, 5 km from Bera de Bidasoa, 150 m, on P. albescens (Hudson) M.
Choisy & Werner, 5 March 1994, J. Etayo 12669. - Spain: Navarra, N of Orbaiceta, N
of Fabrica de Orbaiceta, on Pertusaria sp., 19 July 1991, P. Diederich 9623; Navarra,
Valcarlos, near the French border, 600 m, on P. multipuncta, 19 April 1995, J. Etayo
12922.

This species was known only from the type collection on Pertusaria cf.
ophthalmiza in the British Isles (Hawksworth 1979). Alstrup & Hawksworth
(1990) also mentioned it on Melanelia tominii (Oxner) Essl. (= M. substygia
(Rasanen) Ess.) from Greenland, but the corresponding specimen is likely to
belong to one of the numerous undescribed species of the genus.

Acknowledgments

We would like to thank the curators of K, M, STU and UBC, as well as Drs André
Aptroot and Franz Berger for the loan of specimens, Dr Rosalind Lowen for useful
informations on species of Marchandiomyces and Drs Claude Roux and Dagmar
Triebel for precious comments on the text. The first author is obliged to the DGICYT,
project PB92/0795, for partial financial support.

— 428

References

Alstrup, V. & D. L. Hawksworth, 1990. The lichenicolous fungi of Greenland.
Meddelelser om Grgnland, Bioscience 31: 1-90.

Boqueras, M. & P. Diederich, 1993. New or interesting lichenicolous fungi. 3.
Karsteniomyces llimonae sp. nov. and Sclerococcum serusiauxii sp. nov.
(Deuteromycotina). Mycotaxon 47: 425-431.

Diederich, P., 1990. New or interesting lichenicolous fungi. 1. Species from Luxem-
bourg. Mycotaxon 37: 297-330.

Etayo, J. & J. Blasco Zumeta, 1992. Liquenes epjfitos de zonas dridas. El sabinar de la
Retuerta de Pina (Los Monegros, Espafia). Acta Botanica Malacitana 17: 67-78.

Etayo, J. & P. Diederich, 1995. Lichenicolous fungi from the Western Pyrenees,
France and Spain. I. New species of Deuteromycetes. Jn: Flechten Follmann.
Contributions to Lichenology in Honour of Gerhard Follmann (Ed. F. A. J.
Daniels, M. Schulz & J. Peine), Geobotanical and Phytotaxonomical Study
Group, Botanical Institute, Cologne, p. 205-221.

Hawksworth, D. L., 1977. Taxonomic and biological observations on the genus
Lichenoconium (Sphaeropsidales). Persoonia 9: 159-198.

Hawksworth, D. L., 1979. The lichenicolous Hyphomycetes. Bull. Br. Mus. nat. Hist.
(Bot.) 6: 183-300.

Hawksworth, D. L., 1981. The lichenicolous Coelomycetes. Bull. Br. Mus. nat. Hist.
(Bot.) 9: 1-98.

Hawksworth, D. L., 1990. Notes on British lichenicolous fungi: VI. Notes RBG
Edinb. 46: 391-403.

Kornerup, A. & J. H. Wanscher, 1984. Methuen Handbook of Colour, ed. 3.
Methuen, London, 252 p.

Santesson, R., 1993. The lichens and lichenicolous fungi of Sweden and Norway.
SBT, Lund, 240 p.

Sérusiaux, E. (in press). A preliminary list of foliicolous lichens from Madeira, with
the description of a new genus and two new species. Lichenologist.

Sutton, B. C., 1980. The Coelomycetes. Fungi Imperfecti with Pycnidia, Acervuli and
Stromata. Commonwealth Mycological Institute, Kew, 696 p.

Triebel, D., G. Rambold & J. A. Elix, 1995. A Conspectus of the Genus Phacopsis
(Lecanorales). The Bryologist 98: 71-83.

MYCOTAXON

Volume LX, pp. 429-432 October-December 1996

A new species of Botryobasidium
(Corticiaceae, Aphyllophorales) inhabiting the
interiors of brown rotted logs of ponderosa pine’

MICHAEL J. LARSEN and ALAN E. HARVEY
USDA Forest Service, Intermountain Research Station,
Moscow, ID

ROBERT POWERS
USDA Forest Service, Pacific Southwest Research Station,
Redding, California

MARTIN F. JURGENSEN
School of Forestry and Wood products
Michigan Technological University
Houghton, Michigan

Key words -- Botryobasidium ponderosum, B. medium, B.
subcoronatum, Haplotrichum anamorph, California, ponderosa
pine.

ABSTRACT

Botryobasidium ponderosum is described as a new species associated with a
Haplotrichum anamorph. Basidiomata of this fungus were found in the
interior of brown rotted logs of Pinus ponderosa. The new species is similar
in some respects to B. medium and B. subcoronatum.

INTRODUCTION

In 1993 a multidisciplinary ecosystem study was installed at
the Black's Mountain Experimental Forest (BMEF), within the
Lassen National Forest, in northern California. Forest
conditions at BMEF are typical of the Eastside Pine type (SAF
237; Erye, 1980) that extends from Baja, California, north into
central British Columbia. This ecosystem is characterized by a
cool, dry continental climate. Air temperatures vary between -
9°C and 29°C. Annual precipitation averages 460 mm, falling

1

This article is contribution no. 003 of the Black’s Mountain
Interdisciplinary Research Project, Pacific Southwest Research Station, USDA
Forest Service. It was written and prepared by U.S. Government employees on
official time, and is therefore in the public domain and not subject to
copyright.

430

partly as summer thunderstorms, but mainly as winter snow.
Frost may occur in any month. Natural fire, normally a
periodic event in this forest type, has been reduced
substantially because of fire suppression policies that began
early this century. Ecologically, the area has been typed as
Yellow pine/Purshia tridentata/Idaho Fescue (Alexander, 1993).
Soils are Mollisols derived from Quarternary basaltic flow
rocks.

As a part of this study an investigation into the interior
mycoflora of brown rotted Pinus ponderosa Laws logs was
initiated. These logs were stems of overmature trees that were
toppled by wind at least 50 years previously. Before falling,
they were most likely snags with advanced stages of root and
heart decay triggered initially by Heterobasidion annosum (Fr.)
Bref. Three one meter sections one each, from the top, middle,
and base of fifteen such logs were systematically dismantled
and scrutinized for the occurrence of basidiomata. One
conspicuous basidiomycetous species occurring in the interior
of inspected logs proved to be an undescribed species of
Botryobasidium that we have named B. ponderosum.

Data on microscopic characteristics were obtained from small
portions of a basidiocarp squash-mounted in 5 percent KOH and
Stained with 1 percent Phloxine B or lactophenol cotton blue
(Johansen, 1940). Melzer's reagent (Melzer, 1924) was used to
detect amyloid or dextrinoid structures. Capital letters used
to designate herbaria are those of Holmgren et al. (1981).

Botryobasidium ponderosum M. Lars., sp. nov. Fig. la-g
Etymology--Named for the host substrate Pinus ponderosa Laws.

Basidiomatibus effusis, arachnoideis; hyphis totis septatis
fibulis; basidiis 7.5-10(-12) x 4-5 wm, suburniformibus, 6-8
sterigmatibus; basidiosporis 6.5-7.5(-8) x 2.5-3 mum,
navicularibus; imperfecto statu Haplotricho, conidiophoris
longitudinis indeterminatis, 4-5 wm diam., conidiis singularis
or catenulatis, interdum verticillastris, totis septatis
fibulis, 20-27 x 7-9(-10) wm, fusiformibus vel ovalibus,
hyalinis.

Basidiomata effused, hypochnoid, pale gray to:dull white, ca.
100 wm thick, growth indeterminate; subicular hyphae (6-)7-8(-
10) wm diam., with clamp connections at all septa, hyaline to
faintly citrine, some wall thickening in the more basal larger
diam. hyphae; subhymenial hyphae 4-5-6 um diam.,branching
frequently, hyaline, clamp connections at all septa; basidia
7.5-10(-12) x 4-5 wm, suburniform, 6-8 sterigmate;
basidiospores navicular, 6-7.5(-8) x 2.5-3 wm, hyaline,
negative in Melzer's reagent. Cyanophilous structures absent.

Anamorph -- Haplotrichum. Subicular hyphae (6-)7-8(-10)um
diam., clamp connections at all septa; conidiophore length not

431

determinable, 3.5-4.5 um diam., apparently originating from
subicular and subhymenial hyphae, often fragmenting into
individual cells 35-45 x 3.5-4.5 wm (functioning as conidia?),
clamp connections at all septa, hyaline; conidia blastosporic,
born singly, laterally and apically, or frequently catenulate
and then produced acropetally, sometimes appearing verticillate
at conidiophore apex, clamp connections at all septa, including
conidiophore/conidia points of attachment and between catenate
conidia, 20-27 x 7-9(-10) mum, fusiform to oval, hyaline.

Holotype - USA, California, Blacks Mt. Experimental Forest,
Lassen National Forest, on Pinus ponderosa Laws. collected M.
di. ebarsen; 13, 1V 94, FP’ no... 135882, (CEMR) .+,Additional
specimens examined - USA, California, Black's Mt. Experimental
Forest, Lassen National Forest, on P. ponderosa, collected by
MUG AtSeh), So lV 947 nNOS. he 35862). 13 58es7 ll so8s4 et s358e 5:
i35e¢e@, 135887: 135888, 135889). 135890, 135891 ,.135892 .(CEMR)..

Remarks: Botryobasidium ponderosum is readily diagnosed by the
occurrence of clamp connections at all septa in both teleomorph
and anamorph, size [7.5-10(-12) x 4-5 um) of suburniform
basidia and size [6-7.5(-8) x 2.5-3 um) of navicular
basidiospores. Botryosidium medium J. Erikss. (nomenclatural
type examined: B. and J. Erikss. 2126) is strikingly similar to
B. ponderosum but the former lacks clamp connections on
conidiophores and between conidia (Langer, 1994; Holubova-
Jechova, 1969). Basidiospore size in B. medium is considerably
larger [9-12 x 5-6(-6.5) wm] (see Langer, 1994,, Eriksson, 1958.
Botryobasidium subcoronatum (Hoehn. et Litsch.) Donk is also
Similar, especially with regard to spore size and shape.
However, this species lacks the Haplotrichum anamorphic state
(Langer, 1994; Eriksson and Ryvarden, 1973; Chrisiansen 1960;
Eriksson, 1958; Rogers, 1935). However, Linder (1942) reported
that he was able to trace the connection between conidia of
Oidium curtisii (Berk.) Linder and basidia of B. subcoronatum.
Hallenberg (1978), though, has shown that O. curtisii
[(Haplotrichum curtisii (Berk.) Holub.-Jech.)] is the anamorph
Of B., cuntisia,Hallenb., ‘Rinally,~Taibot (1965) cites. an .O1dium
state for B. subcoronatum and describes, in the key to species
of Botrybasidium, conidia that are fusoid and 15-20 x 6-9um,
and basidiospores 6.5-8.5(-15) x (2-)3-4(-5)um. There is
insufficiant data in Talbot (1965) to draw any conclusions with
regard to what species he really had, but we note the
Similarity of the dimensions of conidia and basidiospores to
those of B. ponderosum.

Apparently, Botryobasidium ponderosum is a widely distributed
and ubiquitous fungus in logs in this area. When comparing
it's occurrence in stands harvested fifty years ago vs. non
harvested stands, there appears to be no discernable
difference. The species appears to be restricted to a narrowly
defined ecological niche.

432

ACKNOWLEDGMENTS

We gratefully acknowledge the field assistance and technical support of
christine Siegle, Bert Spear, and Kendall Kilborn of the USDA Forest
Service, Pacific Southwest Research Station, Redding.

LITERATURE CITED

ALEXANDER, E.B. 1993. Ecological unit inventory. Blacks Mountain
Experimental Forest, California. Unpublished report on file ot the Pacific
Southwest Research Station, Redding, CA. 87p.

CHRISTIANSEN, M.P. 1960. Danish respinate Fungi. II. Homobasidiomycetes.
Dansk Bot. Ark. 19: 57-388.

ERIKSSON, J. 1958. Studies in the Heterobasidiomycetes and
Homobasidiomycetes -- Aphyllophorales of Muddus National Park in North
Sweden. Symb. Bot. Upsal. 16: 1-172.

ERIKSSON, J., and L. RYVARDEN. 1973. The Corticiaceae of North Europe. 2.
Aleurodiscus - Confertobasidium. Fungiflora, Oslo. PP. 59-261.

EYRE, F.H. (ed.) 1980. Forest cover types of the United States and
Canada. Society of American Foresters, Washington, DC. 148p.

HALLENBERG, N. 1978. Wood-Fungi (Corticiaceae, Coniophoraceae,
Lachnocladiaceae, Thelephoraceae) in N. Iran I. Iran. J. Plant Path. 14:
38-87.

HOLMGREN, P. K., N. HOLMGREN, AND L. BARNETT 1990. Index herbariorum.
Part I. The herbaria of the world. Reg. Veget. 120: 1-693.
HOLUBOVA-JECHOVA, V. 1969. New species of the genus Oidium Link ex Fr.
emend Linder. Ceska Mykol. 23: 209-221.

JOHANSEN, D.A. 1940. Plant Microtechnique. McGraw-Hill Book co., New
York-London. 523 pp.

LANGER, G. 1994. Die Gattung Botryobasidium Donk (Corticiaceae,
Basidiomycetes). Bibl. Mycol. 158: 1-459.

LINDER, D.H. 1942. A contribution towards a monograph of the genus Oidium.
Lloydia 5: 165-207.

MELZER, V. 1924. L'ornamentation des spores des Russules. Bull. Soc.
Mycol. France 40: 78-81.

ROGERS, D.P. 1935. Notes on the lower Basidiomycetes. Univ. Iowa Stud.
Ls = 430.

TALBOT, P.H.B. 1965. Studies of Pellucularia and associated genera of
Hymenomycetes. Persoonia 3: 371-406.

a DD d D
7 ING

yp

Fig. la-g. a-Subicular hyphae. b-Subhymenial hyphae. c-Basidia.
d-Basidiospores. e-Conidiophores. f-Fragmented conidiophores.
g-Conidia (From holotype FP135881)

MYCOTAXON

PLANISTROMELLACEAE, A NEW FAMILY IN THE DOTHIDEALES

MARGARET E. BARR

a7) Unverwess, Road | Vvotdney, -pritisn Columbia,
Canada V8L 5G8

SUMMARY

The new family Planistromellaceae is presented
to include taxa having ascostromata, interthecial
tissues, and schizogenously formed, periphysate
ostioles. Members of the family are species of
Flanistronmneila, Planistroma,)Loratospora, ‘the new
genus Eryptio, Microcyclus, and Mycosphaerellopsis.
New combinations are proposed for Planistromella
FUCCI PCR RA eit. acervatca, Pa Parry i. fe hUpL.O ae reo la.
fp pink, and Mierocyelus kawagoel 1°:

Several genera have been described recently as
Members, of ¢Che "Dothideartes (sy istry but ao not :£¢t ) in
any known family. These fungi resemble members of the
Pseudosphaeriaceae in the formation of multiloculate or
uniloculate ascostromata composed of brown, pseudoparen-

ehymatous, celis, in lacking any true peridial structure’,
and in the presence of interthecial tissues surrounding
the basal) layer of “asc. The locules open schizogenously
Dyt a. ‘periphysate “ostiole, “rather “than py. a” simple,

lysigenous pore or by dehiscence of a caplike structure.
ihe “ascli) /ascospores, and anamorphs “are quite similar ‘to
those of species in the Dothideaceae (Mycosphaerellaceae),
bute in’ those’ taxa’ the’ “intertheetal “ tisdues | are’ ‘ess

extensive and the apical pore is formed lysigenously. For
these genera, anda group of other taxa whose ostioles and
tocules sdre' ‘similar, a. new, family must “be: erected,” The

diagnosis of the family follows, with notes on the genera
MOeCeCONSTICULe Lt at) thie. time.

Planistromellaceae M.E. Barr, fam. nov.

Ascostromata multiloculata vel uniloculata immersa
vel erumpentia pseudoparenchymata. Ostiola schizogena
petiphysata.. Asci bitunicati fissitunicati, basali oblongi
vel saccati. Cellula interthecialia. Ascosporae hyalinae
vel fuscae aseptatae vel septatae. Status anamorphosus
pycnidia) vel acervuli vel “~spoerodochia, conidia holo-
blastica hyalina vel fusca aseptata vel septata. Status

434

spermogonicus pycnidia, spermatia phialidica hyalina
aseptata. Genus typicus: Planistromella A.W. Ramaley.
Ascostromata multiloculate or uniloculate, immersed
to erumpent, composed of brown, pseudoparenchymatous
cells;) locules(,small, opening (Dy schazogenously formed,
periphysate ostioles. Asei (bitunieoate,) fissitunieates

oblong. or saccate, in’basal layer, often interspersed with
and overtopped by cellular remnants of interthecial

tis sucsis: Aseospores, (hyaline). oe. sliehely\ pigmented:
yield Vow shi Go brown ds hy, aseptate’ or one. (to! ‘several
transversely septate; wie ae tinder smooth, occasionally
surrounded" by | gel icoatings" contents “guetulate:. Anamorphs
where ) known) forming. pyenmidia | Locules cog’ dace y ulin one
stroma -or bearing “conidta jover, stroma surface, prior, toe
locule formation; conidiogenous cells SinOnet conidio-
genesis holoblastic; conidia hyaline to brown, aseptate or
one to several transversely septate; wall smooth or
verruculose, bearing one or more apical appendages at
times. Spermatial state -developing, an) ) thesesane: com
separate locules; conidiogenesis phialidic; spermatia
mMminute..hyaline)aseptate...\) Biloprophie.. hemi biotrophic. or
Maturing in, dead tCissues. 9 Type genus: “Plenistromelia:

Planistromella. A.W. .Ramaley (Mycotaxon 47: 260.1993) nas
as.) the: type . species, P.. yuceifoliorum AW. opRamaley; {Te
forms biseptate ascospores. The anamorph, Kellermania
yYuccifoliorunm AsW i 'Ramaley’; develops) ‘sin: “the same on
similar stroma as the ascomatal locules, and spermatia may
be produced from phialides in the conidiogenous locule.
In her studies elucidating the diverse fungi that inhabit
leaves of members of the Agavaceae, Ramaley (1993, 1995)
has. -described!) two\c additional ispecies of -Phanistromelia,
each forming uniseptate ascospores, and their anamorphs.
The second species, P. uniseptata A.W. Ramaley, has as
anamorph, Kellermaniavyuccigena 7ellis-& Everh. Theswchdad
species:,..P. torsiftoliorum A’W....Ramaley, has .as: anamorphare
uniseptata (Morgan-Jones, Nag Raj & W.B. Kendr.) Nag Raj.
Three more species are added to the genus, each in leaves
of members of Agavaceae. An, anamoxph. is. linked. circume
stantially <ojvthe Garett), and. second ior these vspecties:,

Planistromella ywucoisena (MM. Ea Bary) Meh! Barr .comb nome

Basionym: ~Wetusteinina .yuceigena ME. (Bary... Comers
Univ, Misch oan wie rb tO toa Pa OW Shoemaker and Babcock
(1987) redescribed and illustrated, this ‘taxon, whichsthey
rightfully, excluded. trom.) Wet tist einina, ul ty. yo L.phowat
suggesting any possible disposition. The species fits
well in Planistromella; the possible anamorph associated
in the same leaves is Kellermania: anomala (Cooke) Hohn,

Planistrome ljhaacer Vata) Babise & bverhob Mei Baw, Compr
TOV ee Om MLL
Basionym: Dtatrype, acervata Elite Go Rverh se ag,

435

My cOdwe C4 2 Rote 2908.88 x, bidisi wanda Everhart .Gleo2)), had
transferred the species to Diaporthe, but Wehmeyer (1933)
excluded it. Barre Glo Dyiplacedi at. lin Myecosphaeretia.,
although the periphysate ostiole negates that disposition.
This tuspectest may) in) fact. represent a species ‘complex:
Ramaley (pers. communication) has: provided, several
interesting observations. Ascospore sizes vary: 15.20, ox

3-4 pm for the New Jersey collection on Yucca filamentosa
Dern ae BAL ¢ NY ee LA L6 te Oo - eotpm efor alcool lection yon. <
elauca Nutt: ) from North Dakota, (Brenckley..F., Dakoté( 3.43.8,
Nis wl O22 Pee erm Lorees: (collection on Yucca spi Crom
Calipverniany'( as Plowrightua (circumscissa | Ttacy’ \&/yEar le,
ROI: amid (ACL Oe) 21 wS'= 29 SOG 2) ie 6 2 ine For sGoLor ado
collections. A Stigmina anamorph develops over immature
stromata of the Colorado specimens, and spermatia form in
shablow scavities ain) the, sitromar This is,.similar, Go sthe
Stuuaelon= inp,wycospheaerella ideightonii OM, “Moreler,, as
described by Sivanesan (1984), where spermatia and conidia
of Stigmina concentrica.(Cooke & Ellis) Deighton. form ,.in
Onmom pthey ‘stroma: in Chath species the ascosponesy are
Smalig, based 275.04 mt and, conldlaware,) naxnrowes, iand
paler» than! those |»Ramaley, found ‘in ~aher) ,coblections;.
AGaLeional | studies. \on.)) these )/ivariiatilons should prove
instructive.

Pilanistromel Lasiparry i, «Gvarls vine Cooke) be Baia, comb.
NOV cee bpis 3 Wee
Basionym: Endothia parryii Farl. in Cooke, Grevillea

pS TiO 2%, 18 8:5:. The gp tungus*)ihad (;beensMasstened, oto
Plowrighntial by, Rehm v(19.08) cand. to _Dothidella’ by \Theissen
and: sydows (1915) . Because of the reddish pigmentation of

stromatic tissues and the periphysate ostioles of locules,
the species has also been described as Hypocrea agaves
MeUD it eReb Uy Winase SOc 4 My coda lrrancea 192 .5292:5 )1903)uy Later
Lranstierred j)\to .Plowrighetasagaves (Maubl..) *Maubl.. , (Bull.
SOCR BMY COL, -Nrancen »2 Sari bas ep OO)... Another name is
Plowriehteias widlpgansonianaxnikel bexgm.;, pilav Myeolsayl2 i ahs 5.
1906. Rehm (1908) and Theissen and Sydow (1915) indicated
that all the names refer to one species, which develops in
margined spots in leaves of species of Agave and Fourcroya
injsoutheastern USAY Puerto Rico, Mexico, and Guatemala.

Planistroma, wA.W..| Ramalern Mycotaxon, 42. 69." 1991) was
erected for P. yuccipenumlAUW...Ramaley, “a species, similar

in stromata and substrate to those of Planostromella. The
stromata are ,usually ‘occupied..in,) succession by . conidio-=
genous locules, wherein the conidia of the anamorph

Piptarthron pluriloculare A.W. Ramaley are holoblastically
formed, by spermatia produced from phialides in the upper
Regions wiots *the, Jocule -omitim separate loeules,” “and by
ascigerous' locules. Thes-ascocpores, hare. unteellular,
Ramatey~(1992,, 1995) hasiudescribed) twoiadditional species
CP rerinsg Pain “ascospore pskze” pandy shape! and. ino c¢heir
anamorphs: Planistroma obtusilunatum A.W. Ramaley and its

436

Figs. 1-11. Members of Planistromellaceae. Leute 28
Planistromella aQernvata’: ea outline of stroma, 2
ascospores. Se... 2... parryi: ) oo, | owt line lof stromay, uy
ascospores. 5-/. Microcyelus, kawagoeiin ) 55.0 Locule; arn
Stroma. OO, eee ca lie as COEpoT ee) : 8-11. Mycosphaerellopsis
myricariae: 8, ascoma in vertical. sections 9, aseusieeOy
ascospores,” lt. conta lume Standard line, =, L350) pm for. 2,

Si SO. Vee code. ane to AM On 2, GANG Diep CPN et BL,

437

Anamorph Pipterthronm i *aniloculiare AL W.. Ramaley and
Pilani e 7 Oma) nolan ae «AY. iW. Ramaley and its anamorph
Piptarthron crassisporum A.W. Ramaley. All three species

develop in leaves of Agavaceae.

Loratospora Kohim., & Volkm!-Kohlme) (Syst. Ascomye. 12: 10.
1903) was, deseribed for 2h. gesteerid: Kohim.. 26 Volkm: =
Fobila. 01993) .a halototerant: “fungus in; culms) cof \vuncus
roemerianus Scheele. From the description and illus-
teaAtrons, Of ‘this: fungus Le lis a icharter member ‘of the new
family, although no anamorph is known.

One sof the ,early:> photographs) |of ajrnfungus) exhibiting
theryieatures sthat. distinguish) ithe whamilly ols / that’ by
Bectseddt Cpe, “f1eyveiGbe “ot Setrrhias.acicovay \bearn.. }
Sigg. (Mycosphaerella dearnessii M.E. Barr 1972). Ts
species is the cause of the brown-spot needle blight of
pines, better known as Lecanosticta needle blight for the
anamorph, Decanosticta’ acicola (Thim.,) Syd. ese eos e
Telative Mycosphaerella) pini: Rostr A) apud Munk. (Scirrhia
pini A. Funk & A.K. Parker), the red-band needle blight or
Dothistroma needle blight with anamorph Dothistroma

Septospora, .(Dorog,. javM.') (Morelets (is) another \iwidespread
species. Because. of the diseases they ‘cause, these
species have been much investigated. The study by Evans

C2964)-,.0n,) Central: Ameriean ‘collections > provides) “many
details onthe history and »morphology of: these, taxa; sas
welld@as on M. ‘¢ibsonii HOC. Evans (netisecn so net) treated
further here). When these two species are removed from
Mycosphaerella, as they must be, a new genus is needed to
accommodate them.

EEUPE LOW Mabe ary) igen snow.

Ascostromata multiloculata vel uniloculata elongata
erumpentia pseudoparenchymatica. LocuLlisviminmimai, sostioli
periphysati. Asicis bitunicati> basali coblongisveliisaceatti
celluli intertheciali crescentia et compressi. Ascosporae
hyalinae didymae oblongae vel fusiformes. Conidia in
acervuli, holoblastica, hyalina vel fusca fusiformia vel
eylindrica). wel -filiformia” septata . Spermatia hyalina
aseptata. opi. Lyphewus: OLigostiroma! acicola ‘Dearn:
Etymology: from the bursting: forth of ascostromata from
the substrate.

Ascostromata multiloculate or uniloculate, elongate,

erumpent from substrate , pseudoparenchymatous, cells
reddish, or, dark , brown, thyphae abundant’ in’ leaf tissues.
Locules’ small, ostioles. schizogenously formed, peri-
physate. (Asci functionally bitunicate, oblong to saccate,
basal, pushing into and compressing interthecial cells.
Ascospores' hyaline, oblong, “hus ad.d. one septate; wall
smooth; minutely guttulate. Anamorphs forming acervuli in
upper regions ‘of stromata. Contdia holoblas¢ie, hyaline

Com eb 20 wn, fUsotuw Gos, Ce bindune eorrn? bit o ray septate.

438
Spermatia phialidic, hyaline,aseptate’.

Eruptio acicola \CDeann’..)) MVE. Barre seomb. *nov.

Basionym: OLigostroma: ecicola Wearn., eMycologra 719s
sa 9) KAP boc 1A as

Eruptio:vpin ge) (Rostr:.) apude) Munk 2 M bewebarre comb enoves
Basionym: Mycosphaerella pini Rostr. apud Munk, Dansk
Bots An ke) ii) go een See

Microcyelus sace., inwsyds &. Poy syduintCAnn ah yecon, (2a Loam
1904) must also be arranged in the Planistromellaceae, for
Theissen and Sydow (1915) and Muller and Sanwal (1954)
described periphyses in the ostioles of the type species,

Mi Wango lense ws Saceus hy eyan te Riv Svar That * fungus - as
biotrophic in leaves of Milletia (Fabaceae) in Africa. A
better knowns species) teem. ftanctoria Clu): Ars iio mes
Mullew& sans! CL962)3> biotrophie onitepecles- of -Baccharse
(Asteraceae) in Central and South America. This species
has an anamorph known as Pazschkeella brasiliensis Syd.
thatr formsipyenidial. cavities. Anvxthe +stroma., The short
eonidiogenous cells) (that «line: the’cavity” form “aseptate:
hyaline conidia holoblastically. Several other species
have been described dn: ‘the .genus, biotrophic: in tropical
and subtropical regions. Gannon et al. (1995) transferred
another: vspecties as “iMicrnoeyclus \porlierizae )}(Rehm)\ePrck.
Cannon, | Carmaran J&/ AG AyRomeroy and (provided a) *key to

South American species.

Another species should be added, from leaves of
Prunus (Rosaceae) in warm temperate regions of Japan.
This’. species was ‘present on Ya) specimen ‘sent, to’ ‘the

University of Massachusetts phanerogamic herbarium, and
the infected leaves were brought to my attention by the
late Harry Ahles. Dr. Kivi Katume to’ kindly furnished \me

with, partiss: of; twoccecokleetions mot this fungus? Pron Syees
with» correction: ot mspelbing for the: specific (epithet? sand
with notes and translations from the, Japanese description.
Additional probable synonyms include: Coccoidea tsurudae
Harayg/fBulloyrShizuokalePreiiin) Agriicay Soc, |) |\(2a7 3 Ai eno oe
Coccoidella tsurudae (Hara), Hara,’ Jubyogaku-kekuron » 82:
1923; amd: \Coceoideila $j spinulosae “(Henn. )> "Haray,) ANippon=
gaikingaku) 1699 i736. The: latter® name. is’ » based wan
Diplodina spinulosae Henuy(Hedwigia 432 145.) L904). ee
conidial fungus that\imay be’ the’ anamorph ‘of Mierocycelus
kawagoeii: iCoccoidélla is a.‘stromatic member ‘of-the
Dothideaceae (Barr 1987) whose ascomatal locules open by a
lysigenous pore.

Microcy clus) jkawagoeii*:)(Hara)) ME. Bary, “combe nev: Figen
5-92
Basionym: Polystomella kawagoii Hara, Bot. Mag. Tokyo
2.9. chiro Mel Ome ae
Additional synonym: Coccoidella kawagoii (Hara) Hara,

439

Jubyogaku-kakuron, 85. 1923.

Ascostromata widely erumpent from central basal
hypostroma, 0.5-1 mm wide, composed of pseudoparenchym-
acouse cells. buown, hyphae. swilthinigleat wtissues;..,.Locules
numerous over entire exposed surface, 65-90 pm wide, 90-
130 «pm. high: .ostiole, periphysate,. ASei.gne biagal )Laye x,
35-70 x 10-12 pm, few pseudoparenchymatous cells remaining
above. Ascospores hyaline, iobovoid 61 2-138620). xed -5 opm;
one septate, slightly supramedian, upper cell shorter and
Wider athan) Lowverce wall smooghies Contents .euttulatges, In
leaves of Prunus. spinulosausiebs 6é4Zacc..,and Pos zippeliana
Miquel., Japan. Material examined: P. spinulosa,Yamaguchi
Pref. Takamizu-mura, Kumage-gum, 10 Feb 1952, I.. Hino ex
YAM 20168 CNN orion Zap e uaa. Maingize ls.) HONS, eer re £7.
Chie si non shat.) 30 Apa.) TORO eH ul Lobe), ire MAS SONY), 35 oP.
zippeliana (immature) , Kyushu, Kagoshima Pref. Mee
Takakuma, 2 Nov 1941, S. Wakabayoshi. ex YAM 20166 (NY).

Ny caspuaerellopsis, Hohn. Cann, Mycol. 16my 1.57 “119 bey.

Otheravtaxa.. that “show, similar | anatomy, ‘and:..form, the
periphysate ostioles schizogenously produce uniloculate
ascomata as immersed biotrophs in leaf tissues. Sphaeria
myrLcariae “Fuckel has» been) assigned. variously, as
Sphaerella myricariae (Fuckel) Sacer GSaccanzdo I tstes 22 ee
Mycosphaerella. myricariae, (Fuckel) Lindau ..(Arx. 1949), or
Didymella myricariae, (Fuckel) Petrak (Petrak 1923, Miller
ann Agee TOG. Share SLOT 2.” heakoson 6 eHhawkswortis 099.3.)0) ox
relegated as a synonym of Pseudosphaerella Hohn. (Clements
and ponear, a1 93 15) 3 THiS kun eis ts phew type sspecies, of) a
Cis ti n.c.cive genus Mycosphaerellopsis that sypresently.
ineludes, only MM .,\myrileariae, (Fuckel) Hohn aGhios. 811.) x
Hohnel (1918) considered it to belong in the Pseudosphaer-
iaceae, because of the abundant interthecial cells
remaining between and above, the .maturing, asci. The
immersed, uniloculate ascostromata in leaves of Myricaria
(Tamaricaceae) in Europe form darkened apices that bear a
few short hyphae, and open by periphysate ostioles. The
OBLONG wetA esaccate “asc. are basal! and “contain, light
yellowish, broadly obovoid, one-septate ascospores, 14-20

aD OF atin An yassociated pyenidial (‘anamorph. is. also
present in, the leaves; the short conidiogenous. :cells are
IO uO Dal 2st. 7c. . producine hyaline ; oblong, uniseptate

Conrad ha.. 722), 0-29 (3005-64, sam

Luetrell. (1973) mentioned three iother taxa that ‘form
ostiolar periphyses and lack pseudoparaphyses. All three
form a hamathecium composed of short periphysoids that
arise from the upper regions of the peridium and extend as
periphyses in the ostiole. Metacapnodium juniperi (W.
Phillips «&) Plow...) Spec.ueis, thenytype votwthe genus, jand -ot
Metacapnodiaceae (Hughes & Corlett in Hughes 1972), in the
Chaetothyriales (Ratio. PLO Ss. Plurisperma dalbergiae
Sivan. is regarded, as ai vspecies.. of /Muellerel la.’ cM.
dalbergiae (CGSalsveleals )) Ngee © Se.) Dyn MUl11. (CURIE Sy) dune 9 eee

440

Verrucariales. Mycosphaerella ascophylli Cotton differs
from typical species of Mycosphaerella in forming both a
periphysate ostiole and narrow periphysoids from the upper
peridium (Kohlmeyer and Kohlmeyer 19/79). These may appear
to be pseudoparaphyses in squash mounts. Lhts\* taxon (iste
widely |) ‘distributed’ mycophycobiont’ in: "which ‘the “algae
Ascophyllum nodosum (L.) Le Jol. and Pelvetia canaliculata
DG.) Se UThuvrecy tareyionever found |) wLehowt. sone orlune us ae
maturity (Webber 1967). AC Ssinilar, Sieuvweton “cers wiher
M. apophlaeae Kohlm. in Kohlmeyer and Demoulin (1981) in
two species of the endemic red alga Apophlaea from New
Zealand (Kohlmeyer and Hawkes 1983). The combination of
characteristics presented indicates that another genus
must be erected, which probably should be arranged in the
Verrucariales.

The genera here assigned to the Planistromellaceae
may be separated by use of the following key.

1. Ascospores aseptate; ascostromata usually multiloc-
ulate; weakly bDLlotrophic (oT ‘saprobic in Agavaceaed.. (nia.
Planistroma

ED AS COS MORES « & Pita Geis ile aus Marc a de eka es aie: ies te cece Tedinngy hetaan CICA hehe cites tin ene eg a
2. Ascospores elongate, three-septate; ascostromata
Uni Locubate in? dead "culms Mob, June U Sm Shite. Loratospora
2; “ASCOSDOL E'S’ ONE OC AEWOe Se pila be. ay i. ae cueie Lie yeaa 3
34) \AsScospores’ broadly) obeveotd; “ascomata, uniloculate an
Livingeor avin. Leaves "on Myr i car ra’). Mycosphaerellopsis
3. Ascospores narrower, oblong to fusoid; ascostromata
UstialPypmuleriocu Pace hie) rel, te tees thle t kw (oie) 2 Maines oem nate acre ae 4
4. Ascostromata widely erumpent, biotrophic in leaves
and stens (of, Va‘toms op Lanes so iin Gin este oes Microcyelus
4. Ascostromata less erumpent, biotrophic, hemibio-
Lropnie. Or) Saproe moped Clade co satue lS ieds oar ellen eer eRe ear neon 5
SAO EROCL Oph Pei Vea Ves Ob 2 BIW ed. oak oe wake bebeniieh eae Eruptio

5S. Blotrophic. hemibiotrophic or saprotrophic, InvAgavaceace
Planistromella

Acknowledgments
Ee tthank Drsv ita skohimeyer “and A.W. Ramaley who.) Kindiry
reviewed the manuscript and offered constructive criticism

in’ the, process:

Literature Cited

Ars, JdecoAy) oon. LO A:9". Beitrage zur Kenntnis der Gattung
Mycosphaerella. Sydowia 3: 28-100.
. Sandy le Mutter’, 97 3) A’ vé-evaluattion “of the

bitunicate Ascomycetes with keys to families and
genera. Stud? Mycol '(Bhaarn) io ole too,

Barim (oMooB: 1972. Preliminary studies on the Dothideales
in temperate North America. Contr. Univ Michigan
Her bia: 23)- 638%,

441

Wy ee LOS 72 Prodromus to Class Loculoascomycetes.
Published by the author, Amherst, MA. 168 p.

Gannon, oP. F.d 6..C. Carmaran Jand Arb" Romero. NASM may
Studies on Ddiotropiive fungi from? Aree n tin ay:
Microcyclus porlieriae, with ja, key,to. Sauth American
species of Microcyeluc a Mycol Res. 99" SS Sos 6.

Releiaie sees Dieta tr Gor De Mena tany einen ta tere SO 2h The North American

Pyrenomycetes. Published by the authors, Newfield,
1 1 fd ne A nat oe
Eriksson? 0) Bh cand) DeLay ihawks worth. 19:9 3.. Outline of
the ) Ascomycetes! = YOO ST a oyec ASCOMY CY "LZ moar e 7.
PVG S 5. at. CO 1984. The genus Mycosphaerella and its

anamorphs Cercoseptoria, Dothistroma and Lecanosticta
on pines.., Mycol.) (Pap., [asc .1-0025

Hohnel, F. von. SEAN Mycologische Fragmente Nrn. 191-
2900 CANN MV COL. L6n Soh ya.

Hughes, S.J. 1972. New Zealand fungi 17. Pleomorphism in
Euantennariaceae and Metacapnodiaceae, two new

families of sooty moulds. New Zealand J. Bot. 10:
225-2462 ¢

Kohlmeyer, df ars) Ewguek © | N/- Demoulin. OSs Panasijtde and
symbiotic fungi on marine algae. Bot. Marina: 24: .9-
rs

rc aa ae » and M.W. Hawkes. 1983. A suspected case of) myco-
phycobiosis between Mycosphaerella apophlaeae (Asco-
mycetes) and Apophlaea spp. (Rhodophyta). J. Phycol.
BO 2a 2 2 OU.

, ands. Kohimeyer 11979 . »Merine Mycology. The

Digher fungi Academie "Presse Ns (690),

eee , and B. Volkmann-Kohlmeyer. 1993. Atrotorquata and
Loratospora: New ascomycete penera (on? Juncus
roemerianus...oyst. Ascomycs 12+. .7.-22.

LOCGreal tbs). L315. 6 LOCKLOaSCOMVCetes., Ch fi. pp. Loo
Zig un. tne, BUngeh,. “VOL. PVA bas. ) Gy OC. L AL NSWOL TE,

F.K. Sparrow and A.S. Sussman. Academic Press, NY.
Mitlers (by Tangs ots Ac vonwA tx: 2h Moyea Die Gattungen der
didymosporen Pyrenomyceten. Beitr. Kryptogamenfl.
Senweigrbh¢ 2). 1-922),
Be ures, 2 A aDG Bos De 1 Sanwa dL. oe Uber die Gattungen
HMicrocycius ‘sacc, | Cocceoidelia wv.) Hohn.., Coceodothis

Theiss. et vsyd.' und weoccodethella.-Theise. “eet Syd.
Sydowla 8: 231-244.

Petr ak ik. BRS AP eae e MyKkologiisches \Notizen. 199), Uber
Didymella, Mycosphaerellopsis und verwandte
Gattungen): Ann. Mycoroe Zbl <3.0..

Ramaley, A. Reeth e FUNGAL GE Yucca, bDacea@ra t..) Fiptarthfon
pluzvitoculare rend | -tta _tekeomorph, Planistroma
yuecigena.* Mycotaxon’. 42; 63-75.

eR to 1 OO 2r. Fungi \from Yueca \Dadcata.. (2. 'Planistroma
obtusilunatum sp. nov., and its anamorph Piptarthron
uniloculare sp. nov. Mycotaxon 45: 449-460.

Pits 1 ey New fungi from Yucca: Planistromella
yuccifoiiorun) igen: GEO SD. novVv., its anamorph,

Kellermania Vico d Pow ox um, sp. FaVOD I yo and

442

Planistromella uniseptata, sp. nov., the teleomorph
of Kellermania yuccigena. Mycotaxon 47: 259-274.
PT ia NS Ae NW New species of Kellermania, .Piptarthron,

Planistroma, and Planistromella from members of the
Agavaceae. Mycotaxon 50: 255-268.

Rehm, dH. 1908. Ascomycetes novi. Ann. Mycods 6") ako
54 a\%

SaCcaragon Pe Aws LSo2.. “SY LLOge (Punpoviie (hace yias oo mol: tae

Shoemaker. RiAgscand biG... Babpeock.. DSS he Wettsteinina.
Cana Gina WB Orn. Mei 19 fon OD

Sivanesan, JA. 1984. The bitunicate Ascomycetes, and their
anamorphs, ° Cramer, Vaduz. /O1'p:

Theissen, F., and H. Sydow. A alesy Die Dothideales. Ann.
Mycol. lS 1ao- 746.

Webber, F.C. MSE Tf Obsierva Eiions sg Onsen eurs Gi eeuie gm liste
history and biology of. Mycosphaerella “ascophny il ia7
PEO Sod Lew MVC Ot no OC wl OU 1 oOo =O Odie

Wehmeyer, L.E. ED BU The genus Diaporthe Nitschke and
Lite vsegregates. Univ, Michigan’ studa (Set wiser 1.91 wii
SHORE.

MYCOTAXON

Volume LX, pp. 443-448 October-December 1996

A NEW SPECIES OF ALYSIDIOPSIS FROM
MEXICO.

Angel Mercado Sierra’, Maria José Figueras’, Julio Mena
Portales’

‘Instituto de Ecologia y Sistematica, Academia de Ciencias de Cuba. AP 8010, Ciudad
de la Habana 10800, Cuba. Facultad de Medicina, Universidad Rovira i Virgili, Carrer
Sant Lloreng 21, 43201 Reus, Espana.

ABSTRACT

Alysidiopsis lignicola spec. nov. isolated from decayed wood in Mexico is
described and illustrated with light and scanning electron microscopy. We provide
a comparative discussion with the other 3 species known inside this genus and
include a key wih all taxa of Alysidiopsis Sutton.

Key words: Alysidiopsis, Alysidiopsis lignicola, Taxonomy, Hyphomycetes,
dead wood, México.

RESUMEN

Se describe e ilustra con microscopia dptica y electronica de barrido Alysidiopsis
lignicola spec. nov. aislada a partir de madera en descomposici6n en Méjico. Se
compara y discute esta especie con las otras tres descritas dentro del género. Se
incluye una clave dicotomica para la clasificacidn de todas las especies de
Alysidiopsis Sutton.

Palabras clave: Alysidiopsis, Alysidiopsis lignicola, Sistematica, Hifomicetes,
madera en descomposicion, México.

INTRODUCTION

During a survey of microscopic fungi collected on leaves and plant debris in
several places of Veracruz State, Mexico, the senior author found an interesting
hyphomycete which colonizes the bark of dead trunks near the sea in the Gulf of
Mexico. From light microscopy and a literature survey we believe we are dealing
with a new anamorph species. Scanning microscopy was employed to obtain
further details of the conidiogenous cells. These studies allow us to classify this
taxon as a new species of Alysidiopsis Sutton (1973). The type species is A.
pipsissewae Sutton. At present this genus includes two other species, A. foliicola
Castaneda & Amoild, and A. yunnanensis L. Guo & X.L. Liu.

The type of this new species is deposited in the Mycological Herbarium of the
Academy of Sciences of Cuba (HACM).

TAXONOMY

Alysidiopsis lignicola Mercado, Figueras et Mena sp. nov. Fig. 16
Coloniae effusae, atrobrunneae, pilosae. Mycelium partim superficiale et partim in
substrato immersum, ex hyphis pallide brunneis, septatis, ramosis, laevibus, 2-2,5
mm crassis compositum. Conidiophora macronematos, recta vel flexuosa, septata,
laevia, brunnea vel atro brunnea, abrupte pallidissima vel fere subhyalina et
irregulariter ramosa ad apicem, 100-200 x 10-15 um ad basim, 5-7 ym crassis
supra basim, 3-4,5 ym ad apicem, parietibus crasis; rami breves parietibus
tenuibus, 4-20 x 2,5-4 ym, terminales vel laterales, pallide brunnei vel subhyalinis,
septati vel aseptati; teretes, cupiformibus vel ampulliformibus. Cellulae
conidiogenae polyblasticae, integratae, terminalesque vel in ramis discretae,
interdum conidia exorientae, denticulatae; dentes validi, lati, aliquante fuscati ad
apicem, 1-1,5 um alti. Conidia in dentium apice disposita, solitaria vel in brevibus
catenis posita, sicca, elipsoidalia, claviformia, citriformia, ovalia vel aliquantum
irregularia, valida cicatrice fusca ad basim notata, rotundata ad apicem,
subhyalina, laevia, transversaliter 0-1 septata, 3-8 x 1,5-2,5 ym.

Holotypus: In cortice emortuis arboris ignotae. Costa Esmeralda, Veracruz,
Mexico, coll. A. Mercado 15 VIII. 1993, (HACM 9111).

Colonies effuse, dark brown, hairy. Mycelium partly superficial and partly
immersed, consisting of smooth, pale brown, septate hyphae 2-2,5 um wide.
Conidiophores macronematous, straight or flexuous, septate, smooth, branched
towards the apex, brownish to dark brown, thick-walled, becoming almost abruptly
subhyaline towards the apex were it ramifies in a complex manner to give rise to
terminal and lateral, short, branches which can produce conidiogenous cells
directly. Conidiophores 100-200 x 10-15 jm (at the base), 5-7 um above the base
and 3-4,5 ym towards the apex. Branches thin-walled, 4-20 x 2.5-4 am of two
types 1) pale brown, cylindrical and septate and 2) subhyaline, aseptate,
ampuiliform or barrel-shaped. Conidiogenous cells polyblastic, integrated and
terminal or discrete on branches, with many denticles, slightly darker at the top

445

where conidia are produced; they can give rise to new conidiogenous cells.
Denticles 1-1.5 jm high. Conidia solitary or in short chains, dry, ellipsoidal,
clavate, limoniform, ovate or slightly irregular, with a dark, pronounced scar at the
base and a rounded apex, subhyaline, smooth, 3-8 x 1.5-2.5 um, septa 0-1.

Holotype: On bark of dead trunk by the sea. Esmeralda Coast, Veracruz, Mexico,
coll. A. Mercado. 15 Aug. 1993, (HACM 9111).

All the species of Alysidiopsis are characterized by possess macronematous
conidiophores irregularly branched near the apex, blastic and denticulate
conidiogenous cells, and usually catenate conidia very variable in shape and
septation.

Fig. 1.- Alysidiopsis lignicola (HACM 9111)

446

cues

5:

?

4. x7500

x6000;

3

x250;

2

lignicola (HACM 9114).

lysidiopsis

Fig. 2-6- A
x1200; 6. x9000

447

The species here described, A. lignicola, is proposed as new because of the
peculiar colour, shape, size, and arrangement of the conidiophore branches and
conidiogenous cells, and also by the shape and size of its conidia. These
characters clearly separate this taxon from the other three known species in this
genus as seen in the taxonomic key provided. A. pipsissewae Sutton, the type
species, collected on peduncular hairs of Chimaphila umbellata var.
occidentalis in Canada (Sutton, 1973), has brown conidiophores only slightly
paler near the apices and measuring 120-170 x 10-12 um at the base, (5-8 um
wide at the apex). Conidia are pale brown, but darker towards the ends, 5-13 x 3.5-
5.5 pm. A. foliicola Castahfeda & G. Amold, collected on fallen leaves of
Calophyllum calaba. in Cuba (Amold & Castafeda, 1986) features short and
narrow conidiophores up to 50 x 4-5 ym (at base) and O-septate mostly cylindrical
to doliiform conidia, brown in the middle and subhyaline at the ends, 5.5-80 x 2-3
ym. More recently Guo & Liu (1992) described A. yunnanensis L. Guo & X.L. Liu,
collected on leaves of two species of Eupatorium, E. odoratum and E.
colestinum L. in China. This taxon is characterized by conidiophores up to 440 x
4.3-7.6 ym and conidia 5.4-26.0 x 5.4-17.3 am with 0-3 longitudinal or oblique
septa.

A further comparative study of the species of Alysidium and Alysidiopsis with
those of Cladosporium complex is necessary since all these genera are very
similar.

KEY TO THE SPECIES OF ALYSIDIOPSIS

Conidia with transverse, longitudinal or oblique septa,
eta Ma NE VID res ea. ac staes Uhivoe ess seueee ehl ty arch tats sage Secoans A. yunnanensis

Conidia with 0-1 transverse septa... cece ccc ceccecceesceeceesccescessecssesseesseeseeessenes 1

1a. Conidiophores short, up to 50 ym long. Conidia always O-septate, unequally
Para Fe U9 Betis aaa a de Ser be Waa AO dS COA NYE Oe A. foliicola
1b. Conidiophores over 100 1m long. Conidia 0-1 septate. 20.0... Z

2a. Conidiophores brown and only slightly paler near the apex, 120-170 x 10-12
pm at the base, (5-8 wde at the apex). Conidia pale brown, darker towards the
f= W0 2 FS nal BLP AG Ms ao kh RU PUREAAE Pers) 1 Fa iC er OP YO a A. pipsissewae
2b. Conidiophores brown becoming almost subhyaline abruptly towards the apex,
100-200 um x 10-15 um at the base and 3.0-4,5 um towards the apex. Conidia
Ssubnyaline: 3-6'X'h:5-2,O7N Ce et I eee A. lignicola

448

ACKNOWLEDGEMENTS

Angel Mercado is grateful to Gabriela Heredia M. Sc. (Instituto de Ecologia,
Xalapa, Mexico) for her aid in collecting this fungus and for providing research
facilities in her laboratory during part of this work. We all thank Dr. Pedro Herrera
(Instituto de Ecologia y Sistematica, Havana, Cuba) for correcting the Latin
diagnoses and Dr. Enrique Descals for revision of the manuscript. This study was
partly supported by Fundaci6 Ciéncia i Salut.

REFERENCES

Amold, G.R.W. & R.F. Castafieda. (1986). Neue Hyphomyzeten-Arten aus Kuba.
Feddes Repert, 97(1-2): 79-88.

Guo, Y.L. & X.J. Liu. (1992). Alysidiopsis yunnanensis, a new species of the
genus Alysidiopsis. Acta Mycol. Sinica, 11(3): 213-215.

Sutton, B.C. (1973). Hyphomycetes from Manitoba and Saskatchewan. Mycol.
Pap., 132: 1-143.

MYCOTAXON

Aone SE SESS Sa IS 5 Sid SE pe Arai eo at Ok = be A 2 alee UR i, eae Me ie
Volume LX, pp. 449-454 October-December 1996

NEW AND INTERESTING HELICOON SPECIES
FROM SPAIN

Samir K. Abdullah’, J. Guarro” and M.J. Figueras”

"Biology Department, College of Science, University of Basrah, Iraq, 7Unitat de Microbiologia,
Facultat de Medicina i Ciéncies de la Salut, Universitat Rovira i Virgili, 43201 Reus, Spain.

ABSTRACT

Helicoon microsporum sp. nov. and H. fuscosporum, a new record for Spain,
found during a survey of aero-aquatic conidial fungi in Northern Catalunya, are
described and illustrated.

RESUMEN

Se describen e ilustran Helicoon microsporum y H. fuscosporum, un nuevo
registro para Espana. Dichas especies fueron aisladas durante un estudio sobre
los hongos conidiales aero-acuaticos realizados en Espafia.

INTRODUCTION

In a survey of aero-aquatic conidial fungi in a static water habitat and streams
surrounded by deciduous and coniferous trees in Northern Catalunya, Spain, two
species, assigned to the genus Helicoon Morgan, were encountered. One is a new
species of Helicoon, the other is H. fuscosporum Linder, a new record for Spain.
The description of the new fungus is based on its growth and sporulation on

450

decaying pine twigs incubated in moist chamber, because attempts in isolating it in
artificial culture media were unsuccessful.

MATERIALS AND METHODS

Samples of leaves and twigs of deciduous and coniferous trees were collected
from a variety of stagnant water habitats and streams in different localities in
Northern Catalunya, Spain. These samples were washed several times with tap
water and then placed on filter papers moistened with water in 15 cm. Petri dishes
and incubated at 15°C under light. Glistening white or coloured propagules with
helicoid shape appeared on the exposed surface within 4 to 6 days.

TAXONOMY

Helicoon microsporum Abdullah, Guarro et Figueras, sp. nov . (Figs. 1-2)

Coloniae in substrato naturali Pini sp., inconspicuae, albae; mycelium superficiale
hyphis hyalinis, ramosis, septatis, 2.5-3.0 ym latis, compositum. Conidiophora
micronematosa vel semimacronematosa, simplicia, raro septata, recta vel flexuosa,
hyalina, laevia, brevia, 5-20 x 3-4 um. Cellulae conidiogenae blasticae, integratae,
determinatae, denticulatae. Conidia solitaria, acrogena, sicca, hyalina, helicoidea,
cum 3-6 spiris (pleraque cum 4-5 spiris), globosa vel subglobosa, 15-17 x 16-20
um. Filamentum conidiale hyalinum, septatum (4-6 septa in unaquaque spira), sine
constrictionibus in septis.

Holotypus: IMI 370113.

Colonies on rotten twig of Pinus sp. inconspicuous, white; mycelium superficial
composed of hyaline, branched, septate hyphae 2.5-3.0 um wide. Conidiophores
micronematous to semimacronematous, simple, rarely branched, straight or
flexuous, hyaline, smooth, short, 5-20 pm long and 3-4 pm wide. Conidiogenous
cells blastic, integrated, determinate, denticulate. Conidia solitary, acrogenous, dry,
hyaline, helicoid, consisting of a filament tightly coiled 3-6 times (mostly 4-5 times)
in three planes forming a globose to subglobose conidium 15-17 x 16-20 jm in
size; conidial filament hyaline, septate (4-6 septa per coil), not constricted at septa.

Specimens examined: IMI 370113 (holotype), FMR 5492 (isotype), on submerged
rotten Pinus sp. twigs. River Aiguamoix, Valle de Aran, Catalunya, S.K.

451

Abdullah, Spain, 11 November, 1995; FMR 5493 on submerged rotten Pinus sp.
twigs, Alt Emporda, Cantallops, Catalunya, Spain, S.K. Abdullah, 7 November,
1995.

Morgan (1892) erected Helicoon based on H. sessile as the type. The genus is
characterized by forming non-proliferating, barrel-shaped conidia from a filament
coiled in three dimensions. Species belonging to Helicoon are commonly reported
from aquatic habitats or in very shady places colonizing a variety of decaying plant
materials (Abdullah & Webster, 1980; Linder, 1929; Eaton & Jones, 1971; Shearer,
1972 and Webster & Descals, 1981). Apart from their usual natural habitat,
Abdullah and Webster (1980b) recorded H. chlamydosporum Abdullah & Webster
and H. fuscosporum Linder from wet soil at a coniferous plantation near a water
reservoir in Devon, Great Britain. Goos et al. (1986), in a review of Helicoon,
accepted 8 species. Matsushima (1993) described H. peruamazonense on rotten
petiole of Palmae from the Amazon River and its tributaries in Peru. More recently,
Van der Aa & Samson (1994) described H. macrosporum on decaying bark of
deciduous tree in a mixed forest in Denmark.

H. microsporum is similar to H. farinosum Linder and H. sessile Morgan in having
white conidia, but in the former they are smaller. The conidia of H. farinosum and of
H. sessile measure 20-27 x 22-37 ym and 20-30 x 33-59 um respectively. Another
difference from Helicoon species forming white conidia is the number of coils per
conidium. Conidia of the present species have a maximum number of 6 coils
(mostly 4-5) while the conidia of H. sessile have 5-16 coils and 5-8 coils in conidia
of H. farinosum. H. chlamydosporum has globose to subglobose conidia (15-18 x
14-17 ym) with 4-5 coils, but can be easily distinguished from the present species
by the fuscous color of its conidia and by the formation of a large number of
chlamydospores in groups forming dark fuscous to black sclerotium-like bodies.
Conidia of the new species could be easily confused with conidia of Helicodendron
japonicum Abdullah (1987). The latter species produces globose to subglobose,
white conidia (18.5-26.5 x 18-22 jm) with 5-8 coils, but can be easily distinguished
by the fact that conidia of H. japonicum proliferate to form crowded chains and
clusters of white conidia.

Helicoon fuscosporum Linder, Ann. Mo. Bot. Gdn. 16: 326 (1929) (Fig. 3)

Colonies on decaying leaves of deciduous trees composed of dark brown, septate
hyphae, 2.5-3 ym thick and numerous erect conidiophores. Colonies up to 1.5 cm
in diam on 0.1% malt extract agar (MEA) in 2 weeks at 15°C, reverse fuscous
black. Aerial mycelium cottony, loose, dark brown, composed of branched septate
light brown hyphae 2.5-3 um wide. Submerged mycelium septate, dark brown 2-2.5
um wide. Conidiophores erect, macronematous, mononematous, thick walled,
fuscous below, dilute fuscous above, arising from the submerged mycelium, simple

452

Figs. 1-2. Helicoon microsporum (FMR 5492). 1. x1100; 2. x3200

453

Fig. 3. Helicoon fuscosporum (FMR 5494). x950.

or branched up to 150 ym in length and 4-5 um wide, mostly bearing more than
one conidium. Conidia solitary, dry, acropleurogenous, fuscous; conidial filament
3.5-4.5 am, coiled 6-13 times in three planes to form ellipsoidal to oval conidia 17-
22 x 25-50 um size.

Specimens examined: FMR 5494, on submerged decaying oak leaves, Alt
Emporda, Cantallops, N. Catalunya, Spain, S.K. Abdullah, 7 November, 1995;
FMR 5495, on submerged decaying Salix leaves, Arriu de Bordius, N. Catalunya,
Spain, S.K. Abdullah, 12 November, 1995; FMR 5496, on submerged decaying
Quercus leaves, Rio Arria dera, Soberpera, N. Catalunya, Spain, S.K. Abdullah, 12
November, 1995.

This is a very common species in the United States and Europe (Linder, 1929;
Beverwijk, 1953; Abdullah & Webster 1980 a,b). It has also been reported from
Japan by Matsushima (1975). The species was detected several times from a
variety of submerged decaying leaves during this study. The present finding

454

represents the first report for the species in Spain. We have obtained several
isolates in pure culture. Most isolates sporulate very well in plates containing 0.2%
malt extract agar incubated at 15°C under light. One isolate (FMR 5494) was found
to produce abundant chlamydospores in culture. In a previous study, Abdullah and
Webster (1980) reported the production of chlamydospores in pure cultures by H.
chlamydosporum and H. fuscosporum.

ACKNOWLEDGMENTS

The authors are indebted to Prof. J. Webster for kindly reviewing the manuscript,
and Dr. J.M. Escola for correcting the latin diagnosis. This work was supported by
the Fundaci6é Ciéncia i Salut.

REFERENCES

Aa, H.A. van der and Samson, R.A. (1994). A new species of Helicoon.
Mycological Research, 98: 74-76. |

Abdullah, S.K. (1987). Two new species of Helicodendron. Nova Hedwigia, 44:
339-343.

Abdullah, S.K. and Webster, J. (1980a). Aquatic and aero-aquatic hyphomycetes
from Ireland. Irish Naturalist Journal, 20: 49-55.

Abdullah, S.K. and Webster, J. (1980b). Occurrence of aero-aquatic fungi in soil.
Transactions of the British Mycological Society, 75: 511-514.

Beverwijk, A.L. Van (1953). Helicosporous hyphomycetes |. Transactions of the
British Mycological Society, 36: 111-124.

Eaton, R.A. and Jones, E.B. G. (1971). The biodeterioration of timber in water
cooling towers. II. Fungi growing on wood in different positions in a water
cooling system. Material und Organismen, 6: 81-92.

Goos, R.D., Abdullah, S.K., Fisher, P.J. and Webster, J. (1986). The anamorph
genus Helicoon. Transactions of the British Mycological Society, 87: 115-
122)

Linder, D.H. (1929). A monograph of the Helicosporous Fungi Imperfecti. Annals
of the Missouri Botanical Garden, 16: 227-388.

Matsushima, T. (1975). lcones Microfungorum a Matsushima Lectorum. pp. 209
Pl. 1-405 published by the author, Kobe, Japan.

Matsushima, T. (1993). Matsushima Mycological Memoirs No. 7. pp. 75 PI. 1-131
published by the author, Kobe, Japan.

Morgan, A.P. (1892). North American Helicosporae. Cincinnati Society of Natural
History Journal, 15: 39-52.

Shearer, C. (1972). Fungi of the Chesapeake Bay and its tributaries III. The
distribution of wood-inhabiting Ascomycetes and Fungi Imperfecti of the

Patuxent River. American Journal of Botany, 59: 961-969.

Webster, J. and Descals, E. (1981). Morphology distribution and ecology of
conidial fungi in freshwater habitats. In Biology of Conidial Fungi I. (ed. G.
T. Cole and B. Kendrick) pp. 295-355. New York and London, Academic
Press.

MYCOTAXON

Volume LX, pp. 455-469 October-December 1996

HYPHOMYCETES FROM STREAMS: NEW TAXA AND NEW
COMBINATION

Ludmila Marvanova
Czech Collection of Microorganisms, Tvrdého 14, 602 00 Brno, Czech Republic
and

Enrique Descals

Instituto Mediterraneo de Estudios Avanzados (UIB-CSIC), c/o Dpto. Biologia
Ambiental, Universitat Illes Balears, 07071 Palma de Mallorca,
Spain

Abstract: Anguillospora fustiformis sp. anam. nov. and Pleuropedium multiseptatum
sp. anam. nov. are described. Tricladium minutum comb. nov. is proposed for
Scorpiosporium minutum.

Key words: new taxa, new combination, Anguillospora, Pleuropedium,
Scorpiosporium, Tricladium.

Anguillospora fustiformis

Apothecia identified as Hymenoscyphus imberbis (Bull.: Fr.) Dennis by M. Svréek
(Prague), were collected on angiosperm twigs partly immersed in water. Multi- and
monoascosporic cultures gave rise to the hyphomycetous anamorph described below:

Anguillospora fustiformis Marvanova et Descals sp. anam. nov. Fig. 1
Etym.: fustis (L.) = stick

Moniliales. Coloniae atrogriseae, modice crescentes, mycelium aerium abundans,
lanosum, funiculosum. Hyphae pallidae vel atrobrunneae cum parietibus crassis,
nonnumquam rugosis. Primordia ascomatum ex hyphis spiraliter contortis vel
irregulariter intricatis. Conidiophora singularia vel aggregata, apicalia, simplicia vel raro
pauce ramosa, septata, cellulis subinflatis. Cellulae conidiogenae subclavatae, usque ad
25 x 2.5-6 um , rarissime cum elongationibus paucis, percurrentibus vel sympodialibus,
apice truncatae vel convexae. Conidia recta vel paulo curvata, obclavata, subcylindrica
vel clavata, raro in parte media attenuata, (50-)80-260(-390) x 6-10(-12) tum, usque ad
25 septata, apice subulata vel obtusa, basi convexa vel obtusa, extensio basalis rara,

456

excentrica; cellulae nonnullae conidiorum subinflatae; conidia vetusta saepe disarticulata.
Dehiscentia schizolytica.

Habitat: Ex apotheciis ad ramos emortuos prope fluminem parvum iacentes, loco
KoSiariska dicto, montibus Little Carpathians, prope oppidum Bratislava, Res publica
Slovaca, isolata. Holotypus: PRM 842929, ex CCM F-17882, isotypus PRM
842928, ex CCM F-17882.

Colony (2% MA) (mono- or multiascosporic) dark grey, growing moderately fast,
reverse black. Aerial mycelium abundant, hairy, funiculose; hyphae either pale and
thin-walled or dark brown and thick-walled, sometimes roughened. Ascomatal initials
frequent in some isolates, terminal or lateral on the pale hyphae, consisting of coiled to
variously entangled hyphal ends. Ripe ascomata not seen. Conidial sporulation aerial or
aquatic and then above, at or below the water level. Conidiophores single or
aggregated, apical or lateral, simple or rarely with one branch, septate, cells sometimes
slightly inflated. Conidiogenous cells subclavate, up to 25 x 2.5-6 um, occasionally with
a percurrent or sympodial elongation, scar flat or convex, often with a nipple in the
centre. Conidia straight to slightly curved, obclavate to nearly cylindrical to clavate,
rarely attenuate near the middle, (50-)80-260(-390) x,6-10 (—12) um, multiseptate, apex
subulate to rounded, base slightly convex or rounded, sometimes with a central nipple,
basal extension rare, excentric; some conidial cells often variously inflated; conidial
outline irregular or wavy due to cell inflations. Secession schizolytic. In older cultures
conidia often disarticulate into units of one to several cells. Germination from any cell;
microcycle conidiation rare, spent secondary conidiophores simulating conidial branches
(fig. 1 L).

Isolates examined: CCM F-17882, CCM F-14984, CCM F-15084, CCM F-15184, CCM
F-15284, CCM F-16084, CCM F-16184, CCM F-16284, CCM F-16484, CCM
F-16584, CCM F-16684, CCM F-16784, all isolated from ascospores shot onto agar;
apothecia (Hymenoscyphus imberbis) collected on twigs near or in a stream in the Little
Carpathian Mountains, Slovakia, in Nov. 1982 and 1984 by L. Marvanova. B247-A-1,
apothecia on a twig near water, Welshbury Wood, Forest of Dean, UK (OS 683157),
May 1981, E.Descals. Single-ascospore isolate.

A.. fustiformis differs from the broad-spored species of Anguillospora Ingold (1942) by
the typically straight, stout conidia with irregularly inflated cells and by the absence of
the percurrent basal extension. The irregular conidial outline is characteristic also of
the simple conidia of Pachycladina mutabilis Marvanova (1988), which are often
straight and considerably overlap with those of A. fustiformis. However, the former have
a typically inflated basal extension or a bulbous base and give rise to whitish colonies on
2% MA. There is also a significant overlap with conidia of the Anguillospora-like
anamorph of Hymenoscyphus malawiensis Fisher et Spooner (1987),
Fig. 1. Anguillospora fustiformis, pure culture. A: developing conidium. B-E: conidio-
phores. H: ascomatal initials. F,G,I-L: conidia. L: secondary conidium formation
(primary conidium with spent conidiogenous cell). All drawings from CCM F-17882
except H: from CCM F-15284. B-E: scale bar a= 20 um, H: scale bar b = 20 um; the
rest: scale bar c = 50 um.

458

where the conidial dimensions are 80-130 x 6-10 um (however, acording to figs. 20-21
the conidial width may reach 14 um), the conidial cells are somewhat inflated and the
colonies are dark grey like in A. fustiformis. Both anamorphs may be differentiated on
the basis of the more regular outline and nearly twice smaller length/width ratio of the
conidia of H. malawiensis.

The Descals' isolate was derived from sole-shaped, c. 13 um long ascospores from a
disc-like, cream-coloured apothecium with 8-spored asci measuring c. 100 x 10 um.

Pleuropedium

The anamorph genus Pleuropedium Marvanova et Iqbal (1973) was described as
monotypic. Its type species, P. tricladioides Marvanova et Iqbal was based on CCM
F-311, which was isolated from the Slovak Republic. However, the protologue also
included characters from HME (=EXR) 2963 from the U.K. Differences in their conidial
size and shape were mentioned, but considered to represent intraspecific variation. The
first author, who prepared the description, had no living material of HME 2963 at her
disposal and presumed that the conidial ontogeny and colony characters were
identical with those of CCM F-311. Further studies of more specimens in pure culture
revealed two groups of isolates, distinctly different in conidiogenesis as well as in colony
characters. These differences seem significant and stable enough to justify recognition of
a second species, which is here segregated from P. tricladioides on the basis of its
multiseptate conidia with typically acute ends, borne on monoblastic (annellidic)
conidiogenous cells and a green pigment usually produced on 2% MA especially when
submerged and/or exposed to daylight. Because the material in HME 2963 is scanty and
poorly preserved, another, more recent British isolate has been chosen as the type.

Pleuropedium multiseptatum Marvanova et Descals sp. anam. nov. Figs. 2-4, 7 A,B
Etym.: multiseptatus (L.)- referring to the densely septate conidial elements.

Moniliales. Coloniae in cultura artificiali pallidae vel pallide griseae, modice crescentes,
mycelium aerium saepe lanosum, funiculosum, mycelium in agaro nonnumquam cum
cellulis inflatis aggregatis. Conidiophora singularia, praecipue apicalia, usque ad 250 x
4-5 um, in parte distali saepe ramificata. Cellulae conidiogenae monoblasticae, apicales,
raro laterales, apice truncatae, determinatae vel cum elongationibus percurrentibus
paucis. Conidia singularia, apicalia, elementa cum apicibus acutis vel acuminatis, densiter
septata; axis rectus, arcuatus vel flexus ad insertionem rami, 38-75 (-100) x 3.5-6.5 um,
typice 6-12 septatus, cum extensione basali percurrente, aciculari, 2.5-12.5 (—22.5) um
longo; rami recti vel paulo arcuati, obclavati, cum insertione valde constricta; ramus
primarius (28-)35-65(-98) x 4-7 um, (3)5-10 septatus, saepe ad cellulam secundam
usque ad quintam axis insertus, nonnumquam flexus apud insertionem rami secundarii;
Fig. 2. Pleuropedium multiseptatum, type. A-C, E: developing conidia (arrows:
probable point of conidial secession; highlighting the monoblastic nature of the
conidiogenous cells). D: spent conidiophore with a percurrent conidiogenous cell. Scale
bar = 25 um.

(cL pt ti
a CB ag ae

oe
B
eo
nH le
AG \
|

D

A
f
J \\
a WWE
Sergei ool
eat
Sort CV
Sg
“Teh
‘ My
i

460

ramus primarius distalis rarus, brevior; ramus secundarius 22-55(-75) x (2.5-)4-6.5 um,
3-9 septatus; ramus tertiarius rarus, 22-40 x 4-5.5 um. Dehiscentia schizolytica.

Habitat: Juncus effusus, in flumine Dart, Bellever Bridge, Dartmoor National
Park, Devon, U.K., Mar. 1975, E. Descals et J. Webster. Holotypus IMI 369970,
isotypi: IMI 369971 et Herb. E. Descals A 318-1-7.

Colony (2 % MA) pale beige or pale grey, growing moderately fast, aerial mycelium
mostly woolly, funiculose; substrate mycelium in some isolates with aggregates of
thin-walled inflated cells; a green extracellular pigment appearing typically in agar after
submergence and/or exposure to daylight. Sporulation underwater. Conidiophores
single, mostly apical, up to 250 x 4-5 um, branches acrotonous. Conidiogenous cells
monoblastic, apical or rarely lateral, determinate or with a few percurrent elongations,
scars flat. Conidia single, apical, elements with acute or acuminate ends, closely septate;
axis straight, arcuate or bent at branch insertion, 38-75(-100) x 3.5-6.5 um, typically
6-12 septate, basal extension typically acicular, percurrent, rarely excentric,
2.5-12.5(-22.5) um long; branches straight or slightly curved, obclavate, primary branch
(28-)35-65(-98) x 4-7 um, (3) 5-10 septate, inserted on the second to fifth cell of the
axis, sometimes bent at the secondary branch insertion; distal primary branch occasional,
shorter than the proximal one; secondary branch (in some isolates often orientated
upwards) 22-55(-75) x (2.5-)4-6.5 uum, 3-9 septate; tertiary branch rare, (in one isolate
relatively frequent), 22-40 x 4-5.5 um. Secession schizolytic.

Other isolates examined: CCM F-46594, from foam from Cheticamp River, c. 10 km
upstream from the mouth of Robert Brook, Cape Breton National Park, Nova Scotia,
Canada, Oct. 1994, L. Marvanova. CCM F-14284, from foam from stream nr. Rejviz,
the Hruby Jesenik Mountains, Czech Republic, May 1984, L. Marvanova. CCM
F-15086, from Juncus conglomeratus decaying stems in a draining canal, nr. the village
Mutna, Bystranska dolina valley, Slovenské Beskydy Mountains, Slovak Republic, May
1986, L. Marvanova. CCM F-13289, from foam from the right tributary of the river
Svratka, near the village Kadov, Czech-Moravian Uplands, Czech Republic, May 1989,
L. Marvanova. A319-1-1, from foam from stream flowing through pastures, on road
from Pentrefoelas to Nebo, Snowdonia National Park, N. Wales, U.K., Mar. 1975, E.
Descals. B137-1-12, from foam from stream flowing through Betula and conifer
woodland, nr. Loch Dubh (Strath Rusdale), Scottish Highlands, U.K., Nov. 1979, E.
Descals.

Conidia of P. multiseptatum may resemble aberrant conidia of Tricladium attenuatum
Iqbal, which, however, have thinner elements, the axis being straight or slightly curved
and attenuate near the branch insertions.

This species seems to be more frequent than P. tricladioides. Records documented by
illustrations or collections seen by us are from: Chile (Burgos & Shearer 1987, fig. 1: 5),
Czech Republic (Kotlaba 1983, det. L. Marvanova), Greenland (Engblom ef al. 1986),
N. Zealand (Aimer & Segedin 1985, fig. 3 v), Norway (Brathen 1984, fig. 2 L),

we we mee ee eee ee ee ee ee ee ee ee wee ee ee ee ee ee ee ee eee eee ee eee reese

Fig 3. Pleuropedium multiseptatum, type. Conidia. A and B are regarded as _ typical.
Scale bar = 25um. .

Fig. 4. Pleuropedium multiseptatum. A-C, E, conidia. D, developing
conidium. F, inflated cells. (A,B: HME 2963; C,D,F: CCM F-14284, E:

CCM F-46594). Scale bar = 20 pm.

463

Sweden (Marvanova & Miiller-Haeckel 1981), U.K. (Ingold 1965, figs. 1 A: 3,10,12,
as ?Ingoldia sp., Ingold 1974, fig. 4: 3, as Gyoerffyella sp., Marvanova & Iqbal 1973,
figs. 1-4).

Because the name Pleuropedium tricladioides is now used in a narrower sense, the
diagnosis is amended as follows:

Pleuropedium tricladioides Marvanova et Iqbal (1973), emend. Marvanova
Figs 5,6,7 C,D

Colony (2 % MA) light buff, slow growing, reaching c. 10 mm diam. after 7 days at
20-22° C, compact, glabrous, aerial mycelium in the middle, yellowish ochraceous,
reverse with a diffusible reddish brown pigment. Substrate mycelium with thin-walled,
inflated cells. Sporulation underwater, with or without aeration. Conidiophores mostly
lateral, simple or branched, up to 60 x 2-3 wm. Conidiogenous cells discrete or
integrated, single or grouped, polyblastic, mostly concurrent with conidia, 10-13 x
2.5-3.5 wm, often denticulate. Conidia in fascicles, closely sequential, elements
obclavate, with subacute ends, up to 5 septate; axis straight to slightly sinuous, 26-51 x
3-5 jm, 3-4(5) septate, base flat or papillate or with a short, conoid extension; primary
branch mostly on the second cell of the axis, 30-42 x 3-5.5 tm, 1-4 septate, distal
primary branch occasional; secondary branch often orientated downwards, 20-35 x
2.5-3.5 um, 0-3 septate; tertiary branch rare. Secession schizolytic.

Isolates examined: CCM F-11087 from foam from Allen Creek, nr. Sackville, New
Brunswick, Canada, Apr. 1987, L. Marvanova. CCM F-311 (ex type) , from foam in a
mountain rivulet nr. the chalet Kamzik, Stary Smokovec, Vysoké Tatry Mountains,
Slovak Republic, Aug. 1968, L. Marvanova. CCM F-11777, from foam in a stream nr.
the village Béla -DomaSov, Hruby Jesenik Mountains, Czech Republic, Dec. 1977, L.
Marvanova. CCM F-10783, from foam from River Teign, Glassy Steps, Dartmoor
National Park, Devon, U.K., Feb. 1983, L. Marvanova. A311-1-3, from foam from
stream flowing through acid moorland, River Dart at Bellever Bridge, Dartmoor
National Park, Devon, U.K., Mar. 1975, E. Descals.

Conidia of P. tricladioides are somewhat similar to the lesser branched conidia with
tapering elements seen in some isolates of Varicosporium elodeae Kegel.

Owing to the hitherto broader concept of P. tricladioides, records without illustrations
or voucher specimens are unreliable. If only those accompanied by drawings are to be
considered in addition to our own, the species seems to be rare: Slovak Republic
(Marvanova & Iqbal 1973), U.K. (Ingold 1979 fig. 48).

Scorpiosporium

Iqbal (1974) distinguished his genus Scorpiosporium from Tricladium Ingold by the
"scorpioid" conidial axis (bent at branch insertions), the broad insertion of laterals and
occasional secondary branching of conidial arms before detachment. However, we see a
continuum between S. minutum and Tricladium species and believe it is premature for

464

generic distinction. A microconidial synanamorph was found by E. Descals in one of his
isolates.

Tricladium minutum (Iqbal) Marvanova et Descals comb. nov. Fig. 8
Basionym: Scorpiosporium minutum Iqbal (1974) Biologia (Lahore) 20:17.
Redescription of the species based on British isolates:

Colony (2% MA) pale brown to dark ochraceous, growing slowly, aerial mycelium
abundant, substrate mycelium with groups of isodiametric inflated cells with brownish
walls. Sporulation underwater. Conidiophores semimacronematous, single, simple, up
to 110 x 2.5-3 tm. Conidiogenous cells integrated, monoblastic, single, apical, typically
percurrent, annellations few. Conidia single, apical, septate, ends subulate to acute, axis
usually bent at branch insertions, often twisted between branches, rarely substraight,
slightly tapered towards both ends, 47-105(-120) x 2.5-4 um, sometimes widened
below branches; base truncate or with a subulate percurrent basal extension up to 20 um
long; branches (1)2(3), diverging in different planes, antrorse and broadly divergent to
perpendicular, or the distal retrorse, 5-10(-15) um apart, insertion broad; proximal
branch 22-57(-85) x 2.5-3 tum, distal branch 17-45 um long. A small percentage of
conidia may form dichotomously branching arms before secession. Secession schizolytic.

Microconidial synanamorph: spermatial, on hyphae. Conidiophores mostly lateral,
single, simple, up to 50 um long. Conidiogenous cells phialidic, single, rarely in pairs,
acrogenous, lageniform, 12-15 x 2.5-3.5 um, collarette cup-shaped, up to 4 um long,
periclinal thickening present. Conidia apical, in chains or groups, short-clavate, aseptate,
2-2.5 x 1-1.5 um, base truncate. Germination not seen.

Isolates examined: A-311-2-21 (= IMI 369972), from culm of Juncus effusus, in stream
flowing through acid moorland, River Dart, Bellewer Bridge, Dartmoor National Park,
Devon, U.K., Mar. 1975, E. Descals. B109-3-5 (= IMI 369973), foam from river
flowing through conifers and grassland, Linn of Dee, nr. Aberdeen, Scotland, Nov.
1979, E. Descals. B125-1-2 (= IMI 369974, with microconidial synanamorph), foam
from waterfall, Silver Bridge, Scottish Highlands, Nov. 1979, E. Descals. CCM
F-16887, foam from a small stream , Ogden Mill Cross, Trans Canada Highway, nr.
Sackville, New Brunswick, Canada, Apr. 1987, L. Marvanova.

Along with the description of S. minutum, three names have been invalidly recombined
(Art. 33.2) in the genus, i.e. Scorpiosporium angulatum (Ingold ) Iqbal (1974), S.
anomalum (Ingold) Iqbal (1.c.) and S. gracile (Ingold) Iqbal (1.c.).

Fig. 5. Pleuropedium tricladioides, type. A: developing conidia. D: spent conidiophore
(note the polyblastic conidiogenous cells). The rest: conidia. Scale bar = 20 pum.

466

Pe F
D
ra
| >
Fig. 6. Pleuropedium tricladioides, E. Descals A311-1-3. A,B, developing
conidia. C,G, spent conidiophores. D-F, H-K, conidia. Scale bar = 20 um.

467

Se:

Fig. 7. A, B, Pleuropedium multiseptatum CCM F-15086. C.D, Pleuropedium
tricladioides, type. Scale bar = 20 tm.

468

Fig. 8.
developm
uum.

Tricladium minutum, E. Descals B125-1-2. A-C, conidial
ent. D,F, conidia. E,G, microconidial synanamorph. Scale bar = 20

469

ACKNOWLEDGEMENTS

E. Descals’ contribution to this paper has been partly supported by the Project "Flora
Micologica Ibérica", DGICYT PB92-0012. The staff at the Limnology laboratories of
the Departament de Biologia Ambiental, Universitat de les Illes Balears are thanked for
providing facilities. Prof. J. Webster (Exeter) kindly reviewed this paper.

REFERENCES

AIMER, R.D. & SEGEDIN, B.P. (1985). Some aquatic hyphomycetes from New
Zealand streams. N.Z. J. Bot. 23: 273-299.

BRATHEN, I. (1984). The aquatic stauroconidial hyphomycetes of Norway with
notes on the Nordic species. Nordic J. Bot. 4: 375-392.

BURGOS, J. & SHEARER, C.A. (1987). Hifomicetes acuaticos en un gradiente
bioclimatico en Chile. Bol. Micol. 3: 101-109.

ENGBLOM, E., LINGDELL, P.-E, MARVANOVA, L. & MULLER-
HAECKEL, A. (1986): Foam spora in running waters of southern Greenland.
Polar Res. 4 N.S.: 47-51.

FISHER, P. J. & SPOONER, B. (1987). Two new ascomycetes from Malawi. Trans.
Br. Mycol. Soc. 88: 47-54.

INGOLD, C.T. (1942). Aquatic hyphomycetes from decaying alder leaves. Trans. Br.
Mycol. Soc. 25: 339-415.

INGOLD, C.T. (1965). Hyphomycete spores from mountain torrents. Trans. Br. Mycol.
Soc. 48: 453-458.

INGOLD, C.T. (1974). Foam spora from Britain. Trans. Br. Mycol. Soc. 63: 487-497.

INGOLD, C.T.(1979). Advances in the study of so-called aquatic hyphomycetes. Am. J.
Bot. 66: 218-226.

IQBAL, S.H. (1974). Scorpiosporium minutum gen. nov. sp. nov., an aquatic
hyphomycete. Biologia (Lahore) 20: 17-21.

KOTLABA, F. (1983). List of fungi found on the excursion during the VII. All-State
Mycol. Conf., Ceské Budéjovice, 13-18 Sept. 1982 (in Czech). Sbor. Jihoées.
Muzea v C. Budéjovicich, Piirod. Védy 23: 37-48.

MARVANOVA, L. (1988). New hyphomycetes from aquatic environments in
Czechoslovakia. Trans. Br. Mycol. Soc. 90: 607-617.

MARVANOVA, L. & IQBAL, S.H. (1973). Pleuropedium tricladioides gen. et sp.
nov. Antonie van Leeuwenhoek 39: 401-408.

MARVANOVA, L. & MULLER-HAECKEL, A. (1981). Water-borne spores in foam in
a subarctic stream system in Sweden. Sydowia 33: 210-220.

Tt ed

3! i

TA

Hi ate
eT

re 7
Lh re 4)

f , ae! fd oe, 6 baa ae ia 4\ ren Vk Mii? ij ye Oe AC AL
ike NE + hue ret Lt, BRO Oy LBA prey a a fal wes

f ‘i ‘ ve a7 ane 7 | 7 an 7

Mh ; Rid ayia)

Ae Nf ate i Ur ETE 9 | ra etd "
a ate Pad TAA | y bY pb Rae ay opts) My

Gp shah

ae: ,
r¢ i

;

4 ‘ a
s ¢ ‘
/ he ee

1
if
“A eG

Di g

MYCOTAXON

poU ie) A UR elidel Ie et ORES a MANO rel) aide Al Sie ee a ade Cs SUP OS jl A
Volume LX, pp. 471-480 October-December 1996

ALLRUS: a system for standard description, identification and classification of
Russulaceae

V. Robert* and B. Buyck**

* Mycothéque de l'Université Catholique de Louvain, 3 Place Croix du Sud, B-1348
Louvain-la-Neuve, Belgium.

** Muséum National d’Histoire Naturelle, Laboratoire de Cryptogamie, 12 Rue
Buffon, F-75010 Paris, France

Abstract

An original software package, called ALLRUS, has been developed to
provide an easier, automatic and objective means to identify, classify and describe
members of the Russulaceae.

The main features of the software, the selection and coding of characters,
and some of the problems encountered in the construction of the reference files are
discussed.

Introduction

Russula is’ the largest and most complex genus in Russulaceae
(Basidiomycotina). The initiative for ALLRUS was taken in 1993 when both
authors agreed to collaborate for the computer-assisted identification of species of
Russula and to provide a means to identify, classify and describe Russulaceae from
around the world. An original software program, called ALLRUS, has been
developed for this purpose. Coding of published descriptions of the members of the
genus began in 1994 and the database now contains 609 descriptions for 486 species
of Russula. In 1995, the reference data file was extended to tropical African
Lactarius (in collaboration with A. Verbeken, Gent University) and the other genera
of Russulaceae.

Selection of characters

The choice of the characters and character states to be considered in
ALLRUS was established in five steps:
1. compile a list of all characters described for species of Russula,
2. select characters that have been described for an appreciable number of taxa,
3. identify the existing diversity in terminology for each selected character,
4. evaluate descriptive terms for similarity and synonymy,

472

5. propose a coding procedure that distinguishes the range of diversity of
terminology with a minimum loss of information.

Apart from the character-set, each description was also given its own
"personal form" that includes the names of genus and species, literature source,
synonymy, other literature references, the date of last revision.

Characters used in the compilation of the reference data file (steps 1 and 2)
were obtained from the description of species of Russula given in Adhikari (1988,
1990), Bills (1984, 1985, 1986, 1989), Bills and Miller (1984), Blum (1962),
Burlingham (1913), Buyck (1988, 1990, 1993, 1994, 1995), Chiu (1945),
Einhellinger (1985), Harkénen et al. (1993), Heim (1938) Hesler (1960, 1961),
Homola and Shaffer (1975), Hongo (1955, 1968), Kauffman (1908), Ktihner (1975),
Marchand (1977), Murrill (1916), Romagnesi (1967), Sarnari (1984, 1991, 1992,
1993), Schaeffer (1952), Shaffer (1962, 1964, 1970a, 1970b, 1972, 1975, 1989,
1990), Singer (1938, 1939, 1950, 1957), Singer et al. (1983), Thiers (1994).

Information offered in descriptions has often been reduced by the creation of
categories or attributes (step 3 and 4). The number of character states or categories
depends on the nature of the character concerned. Simply choosing between presence
or absence of a certain character is often inadequate or impossible. On the other
hand, it is not realistic to include all existing and often ambiguous or confusing
descriptive terms. For example, in the encoded English descriptions of Russula, the
edge of the lamellae is described by the following qualifying terms: entire, smooth,
even, laciniate, sinuate, fimbriate, undate, undulate, eroded, appendiculate, blunt,
dentate, denticulate, crenate, crenulate, serrate, serrulate, lacerate, erose, floccose,
flocculose, fuscidulous, punctate, flattened, depressed, subcanaliculate, canaliculate,
etc. In addition several quantifying attributes were used to modulate these terms:
strongly, slightly, poorly, heavily, largely, smoothly, minutely, entirely, scarcely,
rarely, often, usually, mainly, very, etc. The complexity is even greater if one
compares these terms with a parallel terminology in descriptions written in French,
German or Italian.

It is clear that not only the selection of described characters, but also their
expression or format (similar technique, similar circumstances, similar terminology,
etc.) is essential in establishing complete comparability between descriptions of
characters and thus in making them more accessible for computerization.
Information should be presented in a standardized format and in an unambiguous or
objective - if possible quantitative - way (how many is "numerous" ?). Absence of a
generally accepted terminology and methodology make it often very difficult to
understand exactly what is meant in a description and reduce comparability to a great
extent. One generally has to be acquainted with the adopted terminology and
description habits of all concerned taxonomists (if these are explained at all). This
clearly shows the need for a standardized international descriptive terminology.

Finally, different types of algorithms were developed to code for different
types of characters (step 5) in a way that allowed for the maximum number and
degree of variation of intermediary states without imposing severe restrictions on the
information contained in the reference files (i.e. all measurements in a description
are encoded as such and not reduced to series of categories).

Complex features were described using several sub-characters, e.g. :

- the structure of the pileipellis (4 sub-characters : degree of gelification, presence of
thick-walled elements, presence of inflated cells, orientation of extremities),

473

- the type of vegetation (4 sub-characters : principal biological form, density of
vegetation, tree-type, degree of humidity),

- the ornamentation of basidiospores (7 sub-characters : amyloidity of plage,
localisation of amyloidity on plage, shape of spore ornaments, form at base of
ornaments, height of connections between ornaments, degree of reticulation, height
of spore ornamentation).The precision of coding of complex features was tested by
comparing computer-generated reconstructions derived from encoded values with
illustrations of the same features provided with the original descriptions (Fig. 1).

A selection of similarity coefficients for the identification process are
preferred because probabilistic identification or dichotomous keys may have
misleading consequences when a limited number of characteristics are considered for
identification (Robert et al., 1994; Pankhurst, 1975a). New and more flexible
algorithms to compute similarity coefficients were developed (Robert and Le
Boulengé, 1996). to minimize the lack of coherent information on species and the
existence of uncertain or missing data (Robert and Buyck, 1994; Pankhurst, 1975b).

Standard ——-@ Amyloid ——S» Ornam. ——® Base of —t Connect.

spore plage hemisvh. & ornam. of like crest
weak ornam. 81Ze different
<= 0.5 um

forms

R. olivaceoviolascens drawn Reticulated
by Imler in Romagnesi

Fig. 1. Comparison between the reconstruction by ALLRUS (using coded
information based on the description of Romagnesi (1967)) of a basidiospore of R.
olivaceoviolacens Gillet, and the drawing of Imler in the same publication.

S-comparisons (for size) were designed for quantitative continuous
variables. Minimum, low mean, high mean and maximum values between 0 and
999.99 can be computed. Values provided in descriptions were encoded directly and
can be compared in several ways using up to 10 different algorithms.

S-characters were used for 4 macroscopic features (pileus diameter, stipe
length and width, hymenophore width, thickness of pileus context), 29 microscopical
measures (length and width of spores, basidia, sterigmata, macropleurocystidia,
marginal cells, extremities of pilei- and stipiti-pellis, pileo- and caulo-cystidia;
length-width ratio for spores; width of pseudocystidia, hyphae of subpellis of pileus
and primordial hyphae; thickness of supra- and sub-pellis in pilei- and stipiti-pellis)
and two measures indicating latitude and altitude.

T-comparisons (for triangle) were designed for some discrete ordered or
unordered multistate variables. Codes between 0 and 7 were used with codes 1, 3, 5
and 7 for intermediate or multistate choices and codes 0,2,4 and 6 for different
character state selection.

474

T-characters were employed for 33 macroscopic features (type of basidiome;
surface, dullness, viscosity, pruinosity, venation, zonation, striation, hygrophaneity
and central shape of pileus; pubescence and shape of pileus margin; separability of
pileipellis; type and attachment of hymenophore; spacing, furcation and consistency
of lamellae, presence of lamellulae, colour and type of lamellae edge; presence of
annulus; insertion, consistency, apical colour, basal shape and shape of stipe,
development of subiculum; context thickness; acridity, presence of bitter taste,
presence of latex) and seven characters for macrochemical tests (SV, FeSO4, Gaiac,
Phenol, Anilin, NH4OH, KOH), 45 microscopic features (amyloidity and
localisation of amyloidity on plage, shape and base of spore ornaments, height of
connections between ornaments, degree of reticulation, number of sterigmata; shape,
apical shape, branching, wall thickness, emergence, contents and SV-sensitivity of
macro- and pseudo-pleurocystidia; shape, apical shape, septation and wall thickness
for marginal cells; type of subhymenium and hymenophoral trama; degree of
gelification, presence of thick-walled elements, presence of inflated cells, orientation
of extremities for pileipellis; shape and ramification of terminal elements in pileus;
distribution, SV-sensitivity, metachromatic reaction in cresyl blue and septation of
pileocystidia; presence of extra-and intracellular pigmentation; presence of thick-
walled elements, presence of inflated cells and orientation of extremities for
stipitipellis; shape and ramification of terminal elements in stipe) and for 6
ecological features (principal biological form, vegetation density, tree-type, degree of
humidity, texture, acidity of soil).

C-comparisons (for categories) were designed for discrete ordered or
unordered multistate variables when several fields make up a single character (e.g.
"pileus color" has 9 fields). Codes 0, 1 and 2 were used with code 1 for intermediate
states.
C-comparisons were used to describe taste, smell, colour of pileus (fresh and
in exsiccatum), colour of lamellae, colour of stipe, colour of latex (fresh, dried and
after exposure to air), colour change of context in stipe and elsewhere on basidiome,
abundance of macro- and pseudo-pleurocystidia, maximal height of spore
ornamentation, presence of clamp connections, substratum, climate and continent.

D-comparisons (for degree) were designed for discrete ordered or unordered
and for some continuous variables, such as spore print colour. Values between 0 and
999.99 were coded for this character.

D-comparisons were used for spore print colour using the data from the
colour charts proposed by Romagnesi (1967) and Dagron (1993).

For all comparisons, a code of -1 was used when data were unavailable
(choosing “?”).

Intermediate characters states were interpreted by the program as selecting
for more than one character state.

A number of useful characters were not included in the data files because of
the enormous amount of data that is missing from existing descriptions (see below).
In some cases, it was better to retain different kinds of information on a particular
feature until a satisfactory solution is found in the future.

Missing data

The identification of a specimen using ALLRUS is the result of a more or
less perfect match between its features and those contained in a particular description

475

(or taxon) encoded in the reference data file. When comparing the data set of a
specimen with a data set of a description contained in the reference file, ALLRUS
ignores all characters that are missing in at least one of the two data sets under
comparison. Unless the user restricts the number of reference species that will be
compared for the identification, the character set of the specimen is compared with
every description or species contained in the reference file.

The result of these comparisons is a listing of similarities for all compared
descriptions in descending order. Because the reliability of the identification depends
on the number of characters that were actually compared in each case, ALLRUS
indicates the number of character comparisons that support each similarity. It is
evident that a 95 % similarity resulting from 143 character comparisons is much
more informative than a 95 % similarity based on 32 character comparisons.

Reliable identification is possible on the conditions that :

1. the data-set is complete for as many characters as possible,
characters are described unambiguously or quantitatively. It is possible to replace
vague descriptive terminology with objective, quantitative data. For example,
spore size can be based on statistical measurements and the number of
pleurocystidia can be estimated following Buyck (1991),

3. mycologists come to some agreement on the validity of the chosen character set
and their attributes for a standardized presentation format of the information
contained in descriptions.

The identification of Russula, or any other basidiomycete for that matter, can
hardly be as reliable as it is for yeasts and many micromycetes because the
descriptions of macromycetes contain many more missing or ambiguous data. The
capital problem of any database on macromycetes remains the need of a more
complete data set (Fig. 2-4).

The standardization of character set and formats for the description of taxa
will contribute significantly to the reliability of identifying of species of Russula and
other macromycetes. The use of standardized formats for descriptions will facilitate
the comparison of morphological and anatomical features between species and the
development of a more reliable system of classification.

As far as Russula is concerned, it is not possible to make additional
assumptions on features without new observations of original material or without
adding from other sources (Robert and Buyck, 1995). Our data set is not composite,
meaning that the description of taxa are not composed of a mix of data obtained from
various sources. We only encoded individual descriptions from literature, often
several descriptions for the same species, without combining them into one and the
same description.

476

% missing data: yeasts

OO TO 20 SOU aie SOM NGO 70. S090) LOO

Via ae Laser 4) ale) wid acre

Fig. 2. Percentages of missing data for yeast species. Statistics based on 643
descriptions of species and 94 characters (mostly physiological) from Barnett et al.
(1990) and Kreger-van Rij (1984). Missing in grey color.

% missing data: Penicillium, Eupenicillium
and Talaromyces

0» '10° 20 30) 40" 50):'60'" 70°80 *90 100

57 Fee UR (Guan rat ~~ oem beat == ma gen]

Fig. 3. Percentages of missing data for Penicillium, Eupenicillium and Talaromyces
species based on 150 descriptions and 129 characters from Pitt (1979). Missing in
grey color.

477

% missing data: Russula

OF LOME20577301 401150160 ON SOR 90100

te OO Sar it go Se)

Fig. 4. Percentages of missing data in Russula species based on 609 descriptions and
143 characters. Missing in grey color.

A data file covering the whole range of morphological-anatomical-
physiological features of any genus of macromycetes is an utopia. Waiting for new
examination of all original material and new specimens of every species on which to
base new and “complete” descriptions is not only unrealistic, but will never cover the
whole range of variation for the features of each species. Completing the
descriptions in a practical way will inevitably require composite morphological-
anatomical-physiological data sets for the taxa by assembling data for the involved
species from different sources. Even when performed by a "specialist" of the group
at hand, there remains an inherent danger of mixing data sets that apply to different
taxa. ALLRUS guarantees complete transparency with information gathered from
different sources by giving the possibility to indicate the original source for each
individual feature (or set of features) in the data file ("using "sensu author").

The software

The established reference data file constitutes the basic information used by
the specifically designed identification software, which has been developed for IBM-
PC in Borland C++ and CA-Visual Objects for Windows. Powerful executable
programs (.EXE) assure easy data handling and file management. In addition, CA-
Visual Objects can use the same type of files as dBase. The management of these
files can be done using dBase, FoxPro, Access or MS-Excel making it possible to
also use the same data set for statistical or other purposes. The coding and reading of
data is done by making the appropriate selection from a succession of chosen
submenus. It does not require memorizing codes or character states since all
information is displayed on screen or can be consulted through help-menus. The time
necessary for coding a species description varies from 30 minutes to one hour
depending on the complexity of the description. The coding of a strain is much
quicker.

A similar software, ALLEV, was developed for the identification of yeasts
(Robert et al., 1994) and after a testing period of nearly two years, we can safely state

478

that the reliability of the performed identifications is excellent. The identification
process is almost instantaneous for yeasts and within seconds for Russula.

ALLRUS offers a completely transparent working environment since all of
the information contained in the data files is accessible to the user. Furthermore, the
results of identification can be checked or interpreted on screen or printed out for
every character or species for re-interpretation. This allows a correct interpretation of
the results obtained. ALLRUS also permits the user to define most discriminant
characters, attribute different weights to selected characters, carry out step-by-step
selections on successively narrower groups of species, modify, add or delete
information contained in any of the data files (one is allowed to create additional
reference files), produce ASCII-files for numerical analysis or for statistical
operations with other software, consult in separate windows scanned illustrations of
macroscopic and microscopic features for any Russula at any given moment during
access of the data file or after an identification or search action has been performed,
create a species concept by merging characteristics of several strains, compute
percentage of missing data per character or per species, manage a literature by
adding, deleting, searching or ordering publications by various key words, etc.

Comments, suggestions or criticism on this project may be sent to either one
of the authors of the present paper.

Acknowledgements

We sincerely thank Andrew S. Methven, Timothy J. Baroni and Grégoire L.
Hennebert for their reviews and comments.

Our thanks are also extended to Henri Romagnesi, Annemieke Verbeken,
Wendy Untereiner and Richard P. Korf for discussing character selection and coding
and for correcting the manuscript.

Literature cited

Adhikari, M.K. 1988. The genus Russula from the Kathmandu Valley. In Watanabe,
M. and Malla, S.B. (Eds.), Cryptogams of the Himalayas. vol. 1.

Adhikari, M.K. 1990. The genus Russula from Nepal (II). In Watanabe, M. and S.B.
Malla (Eds.), Cryptogams of the Himalayas. Central and Eastern Nepal. vol 2:
101-112. Tsukuba, Japan: Dept. of Botany, Nat. Sci. Mus. 1990.

Barnett, J.A., Payne, R.W. and Yarrow, D. 1990. Yeasts: characteristics and
identification, ed. 2. Cambridge. 1002 pp.

Bills, G.F. 1984. Southern Appalachian Russula II. Mycotaxon 21: 491-517.

Bills, G.F. 1985. Southern Appalachian Russula Ill. Brittonia 37(4): 360-365.

Bills, G.F. 1986. Russula vesicatoria and Russula angustispora sp. nov. two
confusing species of Russula subsection Lactarioidae. Mycotaxon 26(2): 607-
620.

Bills, G.F. 1989. Southern Appalachian Russula IV. Mycologia 81(1): 57-65.

Bills, G.F., Miller, O.K. 1984. Southern Appalachian Russula 1. Mycologia 76: 975-
1002.

Blum, J. 1962. Les russules. Flore monographique des russules de la France et des
pays voisins. P. Lechevalier, Paris. 229 pp.

Burlingham, G.S. 1913. The Lactarioidae of the Pacific coast. Mycologia 5: 305-
SLL

479

Buyck, B. 1988. Etude microscopique de spécimens-types de Russules tropicales de
la sous-section Diversicolores. Mycotaxon 33: 57-70.

Buyck, B. 1990. Révision du genre Russula en Afrique Centrale. Thése de doctorat,
Rijksuniversiteit Gent. 590 pp.

Buyck, B. 1991. The study of microscopic features in Russula. 2. Sterile elements of
the hymenium. Russulales News 1(2): 62-85.

Buyck, B. 1993. F. Ill. Champignons Afri. Centr. 15: 335-408, pl. 55-68.

Buyck, B. 1994. F. I1l. Champignons Afri. Centr. 16: 409-539, pl. 69-87.

Buyck, B. 1995. F. Ill. Champignons Afri. Centr. 17: 540-000, pl. 88-93. (in press).

Chiu, W-F. 1945. The Russulaceae of Yunnan. Lloydia 8(1): 31-59.

Dagron, Ch. 1993. A new color chart for the precise determination of spore print
colour in Russula. Russulales News 2: 50-51.

Einhellinger, A. 1985. Die Gattung Russula in Bayern. Hoppea, Denkschr. Regensb.
Bot. Gesell. 43. 348 pp.

Hark6nen M., Buyck B., Saarimaéki T. and Mwasumbi L. 1993. Tanzanian
mushrooms and their uses 1. Russula. Karstenia 33: 11-50.

Heim, R. 1938. Les Lactario-Russulé du domaine oriental de Madagascar, un essai
sur la classification et la phylogénie des Astérosporales. Prodrome a une _ flore
mycologique de Madagascar et dépendances, I. Paris. 196 pp.

Hesler, L.R. 1960. A study of Russula type. Mem. Torrey Bot. Club 21: 1-59.

Hesler, L.R. 1961. A study of Russula type. Il. Mycologia 53: 605-625.

Homola, R.L. and Shaffer, R.L. 1975. A new Russula of the subsection Nigricantes
from Northeastern North America. Mycologia 67: 428-434.

Hongo, T. 1955. Notes on Japanese larger fungi (6). Journ. Jap. Bot. Vol. 30 (3): 73-
79.

Hongo, T. 1968. Notulae Mycologicae (7). Mem. Shiga Univ. Nat. Sci. 18: 47-52.

Kauffman, C. H. 1908. Unreported Michigan Fungi for 1908, with a monograph of
the Russulas of the State. Mich. Acad. Sc. 11: 55-91.

Kreger-van Rij, N.J.W. 1984. The yeasts a taxonomic study, third and enlarged ed.
Elsevier Science Publishers, Amsterdam. 1082 pp.

Kiihner, R. 1975. Agaricales de la zone alpine. Genre Russula Pers. ex: S.F. Gray.
Bull. Soc. Mycol. France 91(3): 313-390.

Marchand, A. 1977. Champignons du Nord et du Midi. V. Les Russules. Hachette.
303 pp.

Murrill, W.A. 1916. Illustrations of fungi. - XXIII. Mycologia 8: 121-124.

Pankhurst, R.J. 1975a. Identification by matching. In: R.J. Pankhurst (ed.),
Biological Identification with Computers. Academic Press, London and New
York. 79- 91.

Pankhurst, R.J. 1975b. Identification Methods and the Quality of Taxonomic
Descriptions. In: R.J. Pankhurst (ed.), Biological Identification with Computers.
Academic Press, London and New York. 237-247.

Pitt, J.1. 1979. The genus Penicillium and its teleomorphic states Eupenicillium and
Talaromyces. Academic Press, London. 634 pp.

Robert, V. and Buyck, B. 1994. Missing data in mycology. Abstr. 5th Mycol. Cong.
Vancouver.

Robert, V. and Buyck, B. 1995. Missing data on European Russula. Russulales
News.4.. 91 pp.

Robert, V., de Bien, J.-E., Buyck, B. and Hennebert, G.L. 1994. ALLEV, a new
program for computer-assisted identification of yeasts. Taxon 43: 433-439.

480

Robert, V and E. Le Boulengé. 1996. Some algorithms for identification and
classification purposes. Jn prep.

Romagnesi, H. 1967. Les russules d'Europe et d'Afrique du Nord. Bordas, Paris.
1028 pp.

Sarnari, M. 1984. Genere Russula. Le Plorantinae a spore colorate. Boll. Gruppo
Mic. G. Bresadola 27: 117-124.

Sarnari, M. 1991. Russula nuove o interessanti dell'Italia centrale e mediterranea-
XIV contributo. Boll. Amer. 23(2): 7-17.

Sarnari, M. 1992. Russula nuove o interessanti dell'Italia centrale e mediterranea-
XVI contributo. Boll. Amer. 26(2): 27-32.

Sarnari, M. 1993. Russule nuove o interessanti dell'Italia centrale e mediterranea.
XIX contributo. Rivista Mycol. 36: 37-54.

Schaeffer, J. 1952. Russula-Monographie. Verlag Julius Klinkhardt, Bad Heilbrunn
Obb. 296 pp.

Shaffer, R.L. 1962. The subsection Compactae of Russula. Brittonia 14: 254-284.

Shaffer, R.L. 1964. The subsection Lactarioidae of Russula. Mycologia 56: 202-231.

Shaffer, R.L. 1970a. Cuticular terminology in Russula (Agaricales). Brittonia 22:
230-239.

Shaffer, R.L. 1970b. Notes on the subsection Crassotunicatinae and other species of
Russula. Lloydia 33: 49-96.

Shaffer, R.L. 1972. North American Russula of the subsection Foetentinae.
Mycologia 64: 1008-1053.

Shaffer, R.L. 1975. Some common North American species of Russula
subsection Emeticinae. Beih. Nova Hedwigia 51: 207-229.

Shaffer, R.L. 1989. Four white-capped species of Russula (Russulaceae). Mem. N.Y.
Bot. Grdn 49: 348-354.

Shaffer, R.L. 1990. Notes on the Archaeinae and other Russula. Contr. Univ. Mich.
Herb. 17: 295-306.

Singer, R. 1938. Contribution a l'étude des Russules (1). 3. Quelques Russules
ameéricaines et asiatiques. Bull. Soc. Mycol. France 54: 132-177.

Singer, R. 1939. Contribution a l'étude des Russules (1). 4. Quelques Russules
américaines et asiatiques (suite). Bull. Soc. Mycol. France 55: 226-232.

Singer, R. 1950. Les Russules de l'Argentine. Rev. Mycol. 15: 125-137.

Singer, R. 1957. New and Interesting species of Basidiomycetes V. Sydowia 11: 141-
PH PR

Singer, R., Araujo, I. and Ivory, M.H. 1983. The Ectotrophically Mycorrhizal fungi
of the Neotropical Lowlands, especially Central Amazonia. Beih. Nova
Hedwigia 77: 1-352.

Thiers, H.D. 1994. The subgenus Compactae of Russula in California. Mycol. Helv.
2: 107-120.

MYCOTAXON

ee ARLE i a A ede SSRI Ne) SIRE AE bd, a, sa ee
Volume LX pp. 481-484 October-December 1996

HELICOON SEPTATISSIMUM SP. NOV. , A NEW SPECIES
FROM TIERRA DEL FUEGO (ARGENTINA)

A. M. Godeas* and A. M. Arambarri**

* Departamento de Ciencias Bioldgicas. Facultad de Ciencias Exactas y
Naturales. Universidad de Buenos Aires. 1428. Buenos Aires. Argentina.
** Instituto Spegazzini, Universidad Nacional de La Plata. Calle 53. Nro. 477.
La Plata. 1900 Buenos Aires. Argentina.

ABSTRACT
A previously undiscovered species of a helicosporous Hyphomycetes,
namely Helicoon septatissimum, collected in Tierra del Fuego, Argentina, is
described, illustrated and compared with similar, previously known taxa.
Exsiccata are available at the Spegazzini Institute.

INTRODUCTION
A ten year survey of anamorphic micro fungi growing on wood in Tierra
del Fuego, Argentina, have led to the discovery of previously undescribed
species (Arambarri & Godeas, 1994a,b). In the present contribution an
additional species of Helicoon is reported.

MATERIALS AND METHODS
Wood samples were collected under the surface of slow running
streams. This material was rinsed in tap water and kept damp, but not
submerged, in closed Petri dishes and exposed to light. After 7 days,
glistening spores appeared on the surfaces.
When examined under microscope, these were seen to contain helicoid
spores. The coiled conidium was unsinkable, a bubble of air at the centre of
the coils acting as a flotation device when submerged.

TAXONOMIC PART
Helicoon septatissimum Godeas & Arambarri sp. nov.

Coloniae in naturae inconspicuae. Mycelium plerumque superficialis,
hyphis brunneo-fuscis, septatis, 3-3.8 um diam.; hyphis aereae laevis.
Conidiophora simplicia, erecta, septata, 40-120 x 4.5 um, saepe in apice
recurvata, subhyalina. Cellulae conidiogenae' terminales, blasticae,
denticulatae. Conidia solitaria, siccae, cochleata, cum 9-14 gyris, hyalina vel
pallide brunnea, doliiformis, 70-75 x 40-45 um.

Holotypus: Argentina. Fueguiae. Departamento de Ushuaia. Bahia
Ensenada. Leg. A. Godeas. 3-94, ad ligna putrida Nothofagi betuloides. |n
Herb. LPS 45382 conservatus est.

482

Etimology: with the most septate conidial filament.

Colony on natural substrate inconspicuous. Mycelium, mostly superficial,
composed of dark-brown, septate hyphae, 3-3.8 um diam; aerial hyphae
smooth. Conidiophores, semimacronematous, mononematous, 40-120 x 4.5
um, simple, erect, straight, sometimes curved at the apex, septate,
subhyaline. Conidiogenous cells holoblastic, integrated, terminal,
determinate, denticulate. Conidia, solitary, dry, acrogenous, at first hyaline,
becoming light brown. Conidial filament 8.0 - 10.0 um diam., multiseptate (up
to 30 septa each coil), tightly coiled 9-14 times in three planes to form a
doliiform conidium 70-75x 40-45 um (Fig. 1).

Fig. 1: Helicoon septatissimum;, 1.- Conidiophore and conidia. 2.-Conidia.
Bars 1 and 2 indicate 50um

483

Substrate: On decaying wood of N. betu/oides under water surface in a slow
running stream.
Distribution: Argentina, Tierra del Fuego. Bahia Ensenada.

DISCUSSION

Our species can be placed in the genus Helicoon establised by
Morgan, 1892; who accommodated those helicosporus Hyphomycetes
characterized by the production of non-proliferating, cylindrical, barrel shaped
conidia. The conidia are borne on distinct conidiophores which may be very
short and inconspicuous. No teleomorph of Helicoon is known.

Linder (1929), recognized 7 species for Helicoon whereas Moore
(1955) and Goos (1987), acepted 9 species for the genus.

A revision of the anamorphic genus Helicoon (Goos et al, 1986)
determined 8 species for the genus characterized by the diameter of the
conidium filament, the number of conidial coils, colour, size and shape of the
conidium and conidiophore and conidium size, .

Helicoon septatissimum is included in the table 1 with the diagnostic
characteristics of the species of Helicoon, the substrate where they grow and
the distribution.

The closest species is H. richonis but it has black conidia and the
conidial filament is less septate than H septatissimum.

REFERENCES
Arambarri, A. M. & A. M. Godeas, 1994. Edmundmasonia gamundiae sp. nov.
A new Hyphomycete from Tierra del Fuego (Argentina). Mycotaxon 52:
91-94.
Arambarri, A. M. & A. M. Godeas, 1994. Menispora fuegiana sp. nov. A new
Hyphomycete from Tierra del Fuego (Argentina). Mycotaxon 52:
95-98.
Goos, R. D., 1987. Fungi with a twist: The Helicosporous Hyphomycetes.
Mycologia 79: 1-22.
Goos, R. D., S. K. Abdullah, P. J. Fisher and J. Webster. 1986. The
anamorph genus Helicoon. Trans. Br. Mycol. Soc. 87:115-122.
Linder, D. H., 1929. A monograph of the Helicosporous Fungi Imperfecti. Ann.
Missouri Botanical Garden 16: 227-388.
Moore, R. T., 1955. Index of the Helicosporae. Mycologia 47: 90-103.
Morgan, A. P., 1892. North American Helicosporae. Jour. Cincinnati Soc. Nat.
Hist. 15: 39-52.

ACKNOWLEDGEMENT

The authors gratefully acknoledge Dr. J. E. Wright (U. de Buenos
Aires, Argentina ) for his critical pre-publication rewiew of the manuscript and
help with the latin diagnosis. Also we thank to Dr. J. Webster (U. Exeter,
England) for confirming the new species.

Our research has been supported by a grant from the CONICET
(Consejo Nacional de Investigaciones Cientificas y Técnicas) as part of the
project "Flora Criptogamica de Tierra del Fuego".

484

pue|bu3 “y's'n
eulusbiy

SpUuBLaUJON
‘aoues4 ‘puejGuy ‘epeueo
SpUBLOUJEN ‘puejbuy
spueyeujeN ‘eunuohiy
‘uapems ‘pue|buz ‘y's'n
pue|6uz “y's'n
‘puejeaZz MeN
‘uapams '‘pue|bu ‘y's'n
puejbu

vsn

uonnqiysiq

UOOIIJaH JO Saloads jo DNSUa}OBJeYO jediIoULd :} ajqeL

SBimy:M} ‘Bunp:p ‘jlos:s ‘sanea] :s| ‘(DOOM :pM :ayeQsqns
MO|J9A:A ‘OYUM:M ‘UMOIG WYHII:q] ‘Snoosny:) ‘MojJaA :A sunOjoOgD

“My'S|'PM jepiosdijje GS ¥Xx0S-1 6S-EEX0E-E7 M O[ISSOS ‘H

PM WJOSII/Op S¥X0ZL-O¥ S¥-0¥XSZ-0L qi = winwissyejdes ‘YH

JEAO SG ¥X0S1-0Z 79-0SXZ8-0S q SIUOYSU *H

S| PIOAO =G'E-0'EXOZ1-S ~=—s- OF -SSXTH-0E winjejdasun|d

S| JBA0-jeplosdi|ja 0°S-S'~x00Z- 0S-O2xXSc-Z1 winsodsoosny'H

PM asogo|b-jeplosdilia O'y-S'2XGp-GL — LE-ZZXLZ-0Z WNSOULES 'H

S|'PpM lepiosdijja_—_ $°S-S"y x008- SP-8CXZE-BL ql winoijdija “H

p's‘si‘pm asogojGqns-asoqoj6 ¢'p-S'ex06-S1

SL-SLXZL-vL } wnsodsophweyyo ‘y

PM JBONI}a-PIOAO ~G'G-G'EXOGL-ZE SP-9EXLZ-FL A winjesne *H
(win) azis

aioydoipiuc|

(wn) azis sjiod jo
wnipluod Jequinn

(win) Inojoo
yUOWeyY wnipluo5
winipluod

ajyeysqns edeyus

saiseds

MYCOTAXON

Volume LX, pp. 485-490 October-December 1996

A NEW SPECIES OF LEPTODONTIDIUM
FROM QUERCUS ACUTISSIMA

A. TSUNEDA’', N. MAEKAWA! and R.S. CURRAH?

'Tottori Mycological Institute, Kokoge 211,
Tottori 689-11, Japan
and
*Department of Biological Sciences, University of Alberta,
Edmonton, Alberta T6G 2E9, Canada

ABSTRACT

A new dematiaceous hyphomycete, Leptodontidium quercuum, is
described. This species is characterized by the formation of darkly
pigmented clusters of chlamydospores in Quercus wood and on agar
media. This taxon most closely resembles L. orchidicola in forming
similar chlamydospores but differs in producing colonies which have a
rusty brown cast, in forming large terminal clusters of conidia, and in
being found in wood rather than in living roots.

KEY WORDS: Leptodontidium, Quercus, chlamydospores, taxonomy.
INTRODUCTION
The dematiaceous hyphomycete genus, Leptodontidium de Hoog, is
characterized by gray to black funiculose colonies, hyaline
conidiogenous cells, and small conidia formed more or less sympodially

(de Hoog, 1977; 1979). Six species were described by de Hoog (1977,
1979), treating L. elatius (Magenot) de Hoog as the type, and three other

Paper No. 305. of the Tottori Mycological Institute.

486

species were subsequently added (Rao and de Hoog, 1986; Currah et
al., 1987; Castafieda Ruiz, 1988). We report herein a previously
undescribed species of Leptodontidium from logs of Quercus acutissima
Carruth., that had been used in the cultivation of Shiitake, Lentinula
edodes (Berk.) Pegler, in Japan.

MATERIALS AND METHODS

Three strains of the fungus were studied and deposited in the
Tottori Mycological Institute (TMI): TMI 32391, the ex-type strain,
isolated from wood of Q. acutissima, Kikuchi city, Kumamoto prefecture;
TMI 32392, from wood of Q. acutissima, Saigo-son, Miyazaki pref.; and
TMI 32420, from wood of Q. acutissima, Morotsuka-son, Miyazaki pref.
Subcultures of these strains are in the University of Alberta
Microfungus Collection and Herbarium, UAMH. In addition, two
strains of L. orchidicola Sigler & Currah (UAMH 5422, from roots of
Platanthera hyperborea (L.) Lindl., and UAMH 8151, from roots of
Artemisia norvegica Fries, both collected in Alberta, Canada) were
examined for comparison.

All of these strains were grown on potato dextrose agar (PDA,
Difco, Detroit), corn meal agar (CMA, Difco), or malt extract agar (MEA,
Difco) at 20°C in the dark. Wood samples were hand-sectioned for light
microscopy (LM). Specimens for scanning electron microscopy (SEM)
were prepared using the method described in Tsuneda et al. (1991).

DESCRIPTION

Leptodontidium quercuum Tsuneda, Maekawa & Currah, sp. nov.
Figs. 1-9.
In cortice interiore et alburno exteriore truncorum quercus,
mycelium sterile, inconspicuum alburnum exterius ab eo discoloratum
brunneum videtur. Chlamydosporae forma irregulares, fusce
pigmentatae, crasse tunicatae, praecipue crescentes in vasibus
trachiisque, et ibi texturas scleroticas brunneas vel nigras formantes.
In PDA, coloniae griseae vel rubro-brunneae, cum margine lata, alba,
submersa et cum mycelio aerio coacti-formi olivario vel rubro-brunneo,

post 8 dies propre 20°C ad 63 mm diam attingentes. Propre 25°C

487

incrementum multo tardius et propre 30°C fere abest. Cellulae
conidiogeneae a hyphis indifferentiatis orientes, hyalinae ad pallide
brunneae, lenes, breves, nodo-formes vel longe cylindratae, plerumque
2-3 wm latae et 1-25 wm longae directaeque, tenuiter curvatae vel
flexuosae, septatae vel aseptatae. Conidia sympodialia vel maifeste
phialidia, hyalina, lenia, tenuiter tunicata, subglobosa ad ellipsoidea,
pyriforma, rare breviter stipitata, 2.44 x 2.2-3.2 um, glomus ter minale
e duobus vel ad 20 vel pluribus formantia. Chlamydosporae crassiter
tunicatae, irregulariter formatae, aggregationes densas vel strata
texturae scleroticae formantes. In CMA pigmentatio tarda. Holotypus:
colonia exsiccata ex TMI 32391, ex ligno Quercus acutissima
Japoniensis, TMI.

ETYMOLOGY: Occurring on Quercus.

Mycelium in the natural substrate inconspicuous, spreading in the
inner bark and outer sapwood regions, changing the color of sapwood to
grayish brown. Conidia not found in the natural substrate.
Chlamydospores, irregular in shape, darkly pigmented, thick-walled,
occurring primarily in vessels and tracheids (Fig. 1), often becoming tightly
packed to form brown to black sclerotic tissues. Colonies on PDA
attaining 51-63 mm diam at 20°C in 8 days (varying with the strain; daily
growth rate between 5th and 8th days = 3.9-4.6 mm/day), much slower at
25°C, almost no growth at 30°C, grayish or reddish brown, margin broad,
submerged, white (Fig. 2); aerial mycelium, felted, initially white becoming
Olive to reddish brown with age. Reverse black when aged. Hyphae
septate, often producing brown exudates (Fig. 3), submerged hyphae olive
to dark brown. Conidiogenous cells, arising from undifferentiated hyphae,
hyaline to pale brown, smooth, short knob-like (Fig. 4) to long cylindrical
(Fig.7) often slightly tapered at the tip, mostly 2-3 um thick and 1-25 um
long, straight, slightly bent or sometimes flexuous (Fig. 5), with or without
thin septa. Conidiogenesis either sympodial (Figs. 4-6) or apparently
phialidic (Fig. 7). Conidia hyaline, smooth, thin-walled, subglobose to
ellipsoid, pyriform, rarely short-stipitate, 2.44 x 2.2-3.2 wm, forming a
terminal cluster of a few to 20 or more conidia (Figs. 4-7).
Chlamydospores thick-walled, irregularly shaped, forming aggregated
masses (Figs. 8, 9) or a dense layer of sclerotic tissue. On CMA
pigmentation of colonies and chlamydospore production retarded.

488

Holotype: a dried colony on PDA from TMI 32391, from wood of
Quercus acutissima from Japan, deposited in TMI.

DISCUSSION

Leptodontidium quercuum and L. orchidicola are unique in the
genus in producing chlamydospores that can form sclerotic clusters of
cells on solid media and intracellularly in plant host tissues. In L.
quercuum, the sclerotic tissue develops and forms a narrow, dark
demarcation line in wood apparently in response to the approach of the
mycelium of Lentinula edodes. Similar aggregations develop in root
cortical cells infected by L. orchidicola (Currah et al., 1987). Sclerotic
tissue develops in agar culture of both L. quercuum and L. orchidicola as
colonies age. Chlamydospores are also formed by L. irregulare (de
Hoog) de Hoog but these are ellipsoidal, reddish brown and solitary or
in short chains (de Hoog, 1977). Leptodontidium irregulare, like L.
quercuum, is known from wood but the morphology and disposition of
the mycelium in or among xylary elements is unknown.

Leptodontidium quercuum and L. orchidicola differ substantially
with respect to their natural habitat and the pattern in which conidia
are borne on conidiogenous cells. The latter is known only from roots of
a wide range of plants native to cool temperate regions where soils are
rich in organic debris. Our isolates of L. quercuum are all from wood.
Conidiogenesis in L. orchidicola involves the production of solitary
lateral conidia measuring 1-3 x 3-6 wm on conidiogenous hyphae.
Occasionally a few conidia will develop sympodially in clusters at the

Fig. 1. Transverse section of outer sap wood of Quercus acutissima
containing sclerotic tissues of Leptodontidium quercuum in vessels
(arrows) and tracheids. Sclerotic tissues become black when aged. Fig.
2. Colony of the ex-type strain (TMI 32391) on PDA, 2 weeks at 20°C.
Fig. 3. Hyphae surrounded with exudates (arrows). Figs. 4-7.
Sympodial (4-6) or apparently phialidic (7) conidia formed on
conidiogenous cells of different length. PDA, 3 months at 20°C. Figs. 8
and 9. Masses of chlamydospores developed on MEA and their enlarged
view. 3 months at 20°C. Bars = 200 um in Fig. 1; 300 wm in Fig. 8; 10
um in Figs. 3-7 and 9.

—

490

apex of conidiogenous cells. The process in L. quercuum results in the
formation of few- to many-spored clusters of conidia, 2.44 x 2.2-3.2
um, at the apex of conidiogenous cells. Conidia develop sympodially
although when large clusters of conidia are observed they appear to
have originated phialidically. The mode of origin of these larger clusters
has yet to be clarified. Unlike L. orchidicola, laterally borne conidia are
rare. Leptodontidium orchidicola generally sporulates readily in culture
although conidia may be few and inconspicuous, whereas L. quercuum
requires several months incubation before conidia will appear.

Acknowledgements: We thank Dr. Michael Hertwig, University of
Alberta, for kindly providing the latin description, and Dr. D. Malloch,
University of Toronto, for reviewing the manuscript.

LITERATURE CITED

Castaneda Ruiz, R.F. 1988. Fungi Cubenses III. Publ. Inst. Invs. Fund.
Agric. Trop. “Alejandro de Humboldt,” pp. 27.

Currah, R.S., L. Sigler, and S. Hambleton. 1987. New records and new
taxa of fungi from the mycorrhizae of terrestrial orchids of Alberta.
Can. J. Bot. 65: 2473-2482.

De Hoog, G. S. 1977. Rhinocladiella and allied genera. In Studies in
mycology. No. 15. Centraalbureau voor Schimmelcultures, Baarn.
pp. 140.

De Hoog, G. S. 1979. Nomenclatural notes on some black-yeast like
hyphomycetes. Taxon 28: 347-348.

Fernando, A.A. and R.S. Currah. 1995. Leptodontidium orchidicola
(Mycelium radicis atrovirens complex): aspects of its conidiogenesis
and ecology. Mycotaxon 54: 287-294.

Rao, V. and G.S. de Hoog. 1986. New or critical Hyphomycetes from
India. In Studies in mycology. No. 28. Centraalbureau voor
Schimmelcultures, Baarn. pp. 84.

Tsuneda, A., N. Maekawa, I. Ohira, and I. Furukawa. 1991. Incipient
decay of Quercus serrata sapwood and its inhibition by an
antagonistic hyphomycete, Leptodontidium elatius. Can. J. Bot.
69: 2797-2805.

MYCOTAXON
ERT pp 491 498 TTR OR hes

BOOK REVIEWS
D. S. Hibbett, Book Review Editor

Ainsworth & Bisby's Dictionary of the Fungi, Eighth
Edition, edited by D. L. Hawksworth, P. M. Kirk, B. C. Sutton,
and D. N. Pegler. 250 x 175 mm, 632 pp, 1995. CAB

International, Marketing Department, Wallingford, Oxon OX10
8DE, UK. Fax: (01491) 833508, e-mail: cabi@cabi.org. ISBN 0-
85198-885-7. US$ 49.95.

The 8th edition of the Dictionary of the Fungi is
everything the editors hope for (p. vii), "A marvelously
imperfect work needed by all." The core work is indeed a
marvel of a dictionary. For over a half-century the editorial
format of the Dictionary has been similar to the definition of
the term in Webster's New World Dictionary of the
American Language: "A record of generally accepted
meanings, pronunciations, etc. which the words listed in it
have acquired up to the time of its publication." The historical
mission of the Dictionary has been to provide logically
indexed access to mycological literature. The editions under
the editorial guidance of G. C. Ainsworth and G. R. Bisby stand
for the most part as reputable, authoritative, and reliable
resources for the mycological expert, the general user and the
public at large.

At best the 8th edition continues to be a thorough record
of terms in mycological literature that spans over 200 years.
Mycological specialists from all over the world will be able to
critically read entries reflecting both familiar and unfamiliar
topics. However, the 8th edition will be accepted uncritically
by the nonspecialist user and especially by the general public.

As the editors predict, the 8th edition is imperfect. The
use of terminology is inconsistent. The taxonomy is
misleading. The touted hypothesis-testing skepticism (paxil) is
especially needed for features such as the classification
scheme, the "accepted" taxa, and the "longer entries." More in
the fashion of an encyclopedia than a dictionary, the 8th
edition contains a noticeable increase in the interpretations of
mycological concepts from the viewpoint of the editorial staff.

Two ascomycete examples are illustrative of
imperfection. A boasted innovation is the abandonment of
"the deuteromycetes as a formal systematic category and [the

492

editors endeavor] to incorporate them in the system of
ascomycetes and basidiomycetes...(p. vii)." The 8th edition
makes a bold attempt to clarify the ideological muddle
surrounding the terminology of Article 59 of the
International Code of Botanical Nomenclature that was
contrived for naming pleomorphic species. Nevertheless, the
definitions of terminology for pleomorphic species found in
Introductory Mycology, 4th edition by Alexopoulos,
Mims, and Blackwell (1996; see review in Mycotaxon vol. 58,
1996) are more precise and useful than those provided in the
Dictionary. The 8th edition definition of the fungal
holomorph is adapted from the seminal paper of G. Hennebert
and L. Weresub in 1977 on nomenclatural terminology, but the
later clarification of these terms by Hennebert in The Fungal
Holomorph is ignored. The term "mitosporic fungus" is
introduced as a biological term for the asexual species "to
replace the form taxon." Confusing exceptions in terms and
definitions are made for the mitosporic species as a holomorph
and the asexual morph of a pleomorphic species. The term
anamorph is given new meaning: Phoma is cited as an
"anamorphic Pleosporaceae," Ascochyta as an "anamorphic"
Dothideales, Mycosphaerellaceae and Leptosphaeriaceae. “Yet
the mitosporic, i.e. nonpleomorphic, taxa of the same name are
not incorporated in the touted "system adapted for the
ascomycetes developed in collaboration with O. E. Eriksson."
Purely mitosporic genera such as Septonema are listed along
with anamorphic genera such as Phoma and Ascochyta, as
designated by the Dictionary, in "a non-systematic category
of genera listed alphabetically" (pp. 41, 594).

The authors expound that "Classifications are
hypotheses that need a period of testing before they can be
commended for general use" (p. vii). The 8th edition is replete
with unexplained modification of taxon definition and
acceptance and an error laden "hypothetical" classification
scheme. The Capnodiaceae is an example of what could be
taken as revisionary taxonomy. The family definition (p. 74)
ignores the periphysoids described from the type species of
Capnodium and erroneously cites a paper with this listing of
this term that is unrelated to the genus. The description
specifies brown ascospores, but the list of genera (p. 545)
includes Scorias and Trichomerium which have hyaline
spores. Why are characters recognized in the literature
ignored in the 8th edition definition? Several genera such as
Anopeltis and Callebaea are are included in the family list;
their individual listing is given as the Capnodiacae with a
question mark. What is the basis of this familial revision?
What is recommended in the primary literature?
Trichomerium is listed with 23 species but the cited recent

493

monograph of that taxon recognized one species. What is the
basis for the revision of this genus? Polychaeton is a final
example. This genus was lectotypified by Hughes in 1976 as a
name for pycnidial species. The genus is deemed in the 8th
edition as a "nom. dub., ? Capnodiaceae." Furthermore, the
anamorphic name "Polychaeton nom. dub." is listed in the
"Systematic Arrangement" under Dothideales, Capnodiaceae!

The classification of basidiomycetes in the 8th edition of
the Dictionary is similar to that presented in the 6th and 7th
editions. The merging of the Gasteromycetes with the
Hymenomycetes is a major improvement over earlier editions.
The 8th edition is a step backwards, however, with respect to
the classification of heterobasidiomycetes. If one relied only
on the Dictionary, one might think that there has been no
progress toward defining natural relationships of
heterobasidiomycetes in the last quarter century. This, of
course, is not true. The key to the classes, subclasses and
orders of Basidiomycota demonstrates why the taxonomy is
flawed; the characters which have been given the greatest
taxonomic importance include basidium morphology (sensu
Talbot) and septal ultrastructure.

"By accepting the distinction made by Talbot (1968)
between 'primary' and '‘adventitious' septa two subclasses may
be distinguished" (p. 47). The familiar taxonomy based on this
differentiation is retained in the subclass taxa
Phragmobasidiomycetidae and Holobasidiomycetidae. However,
despite affirming the taxonomic value of this distinction
verbatim from the 7th edition (1971), the 8th edition uses this
character inconsistently in the resulting taxonomy. The
reason for this inconsistent use is that basidial septation has
not withstood the test of time as a high level taxonomic
character. An example is the family Chionosphaeraceae.
Chionosphaera and Fibulostilbum do not have phragmobasidia
Whereas Stilbum does. It is ironic that a taxonomy based
largely on this character would be commended for general use.

Septal ultrastructure is also used to define the higher
taxonomic groups of basidiomycetes recognized by the
Dictionary. Biochemical characters have largely confirmed
the value of septal characters for use in higher level
Systematics. However, numerous, major errors in
interpretation of septal ultrastructure have resulted in a
nonsensical taxonomy. R. T. Moore was the first to propose that
the simple septum of the Uredinales is in fact a reduced
dolipore septum. His reinterpretation of the rust septum
presumably was sparked by early phylogenetic work using 5S
tRNA sequences obtained from so-called rust yeast cultures. In
these analyses, the rust fungi grouped with the dolipore-
septate hymenomycetes sensu Swann (agarics, polypores,

494

Tremellales, etc.). Rust fungi, however, are not known to
produce a yeast phase, and it has been known for some time
that the yeasts in these studies were actually tremelloid yeasts.
Nevertheless, the 8th edition follows Moore's misinterpretation
of the ultrastructural morphology of the rust septum, and
unfortunately compounds this error by expanding the range
of organisms which are said to have this dolipore septum,
including the Septobasidiales, Agaricostilbales, Atractiellales
(sensu lato), and Heterogastridiales. To make matters worse,
the latter three orders are said to have parenthosomes
associated with these dolipore septa. This is incorrect. None of
the organisms in these orders, nor those in the Uredinales,
have a dolipore septum, reduced or otherwise.

The Dictionary claims that "the analysis underlying
the subdivisions used...is pragmatic," and is geared toward the
"special interests of many of those who consult this
Dictionary." This statement suggests that the taxonomy is not
meant to be a scientifically sound hypothesis, but rather is
designed for ease of use by non-specialists. However, the
touted pragmatism is difficult to perceive when the key is
consulted. As mentioned above, certain characters cited for
particular groups of heterobasidiomycetes are factually
incorrect. In other cases the key shows internal conflict of
characters, as well as conflict with characters cited in the
entries for the taxa. An example is Septobasidiales, which is
said to be in the class Teliomycetes. Some of the characters said
to define this group are karyogamy by spermatia, dolipore
septa, and absence of a basidiome (p. 519). Later in the key,
however, the order is differentiated from the Uredinales by
the formation of a corticioid basidiome (p. 529). It is also
clearly stated in the entry under Septobasidiales (p. 414) that
basidiomata are formed by some members of this group.

Persons consulting the taxonomy presented in the
Dictionary should keep in mind that natural relationships
are not necessarily reflected by this classification, nor were
they intended to be. The three classes of the Basidiomycota and
two subclasses of the Basidiomycetes are polyphyletic. At least
five of the heterobasidiomycetous orders presented are
polyphyletic. The orders Platygloeales and Sporidiales are
highly polyphyletic. The Ustilaginales (-aceae) is
polyphyletic due to the inclusion of Microbotryum. The orders
Atractiellales, and Tremellales have been modified from the
most recent treatments of these taxa.

The mini-essays called "longer entries" provide concise
summaries of diverse current topics of interest to mycologists,
but they should be separated from the terms and their
definitions. Here the Dictionary has taken on the aspect of
an encyclopedia. These longer entries may encourage

495

editorial license in the standard entries.

In summary, the tremendous effort necessary for the
compilation and indexing of new information in the 8th
edition is to be commended. The logically indexed definitions
of terms, with validating references to the primary literature,
should be continued. Editorial opinion, however, should be
confined to the preface and the foreword. Imposition of a
Dictionary-endorsed de novo classification on the taxonomic
entries masks the diversity of taxonomic opinion. The effort to
expand the mission of this work with the incorporation of
interpretative revisions of terms and concepts appearing for
the first time in the Dictionary must be viewed with
Skepticism in preparation for the 9th edition. Don R. Reynolds,
Natural History Museum, 900 Exposition Boulevard, Los
Angeles, CA 90007, and Eric Swann, Department of Plant
Biolology, University of Minnesota, St. Paul, MN 55108-1095.

Polyporus (Basidiomycotina) and Related Genera, by
Maria Nufiez and Leif Ryvarden, with drawings by Leif
Ryvarden. 210 x 155 mm, 85 pp, 1995. Synopsis Fungorum vol.
10. Fungiflora A/S, P.O. Box 95, Blindern, N-0314, Oslo, Norway.
Fax: 47-2285-6717. ISBN 82-90724-15-2. NOK 100.

This is a world-wide monograph on the genus Polyporus,
type of the family Polyporaceae sensu Donk (Aphyllophorales,
Basidiomycotina), and of its morphologically related genera
Echinochaete, Pseudofavolus, and Laccocephalum. The book is
divided in the following nine sections: Introduction (which
presents a taxonomic and nomenclatural history of the genus
Polyporus, its typification, and the species concept adopted by
the authors), Taxonomic characters (a detailed explanation of
the characters found in the genus together with a discussion
on the relative taxonomic weight of each character), Ecology
of the species and infrageneric groups, Genera related to
Polyporus, Key to genera and species, Acknowledgments,
Excluded species, References, and Index.

This manual sets the present knowledge on the treated
genera and is a useful and indispensable guide to students of
polypores. The taxa are presented alphabetically, their
descriptions being complete and according to a standardized set
of characters, including references to cultural studies. This is
most useful to the reader because of the clarity of the
presentation and the ease of comparison with other species.
Drawings, however, are scanty, restricted to spores and to the
setoid elements present in Echinochaete. One would appreciate
having figures of the vegetative hyphae that, according to the
reviewer's experience and the literature, present variation

496

within the genus Polyporus.

This study is based mainly on morphological characters.
In spite of the authors' claim that "the biological species
concept has been the criterion used in this work," neither
experimental data on mating tests, nuclear behavior of the
mycelium, isoenzymological, nor DNA studies are presented.
Nevertheless, other non-morphological key features of
Aphyllophorales taxonomy, such as the type of rot produced by
these organisms, have certainly been considered. Indeed, the
restoration of the genus Laccocephalum, based on its
association with a brown rot but long included as a synonym of
Polyporus, a white rot genus, is one of the main achievements
of the work.

Several species complexes have been treated as such,
with no attempt to resolve them. Thus, they have been
accepted in a very broad sense. This is especially so with
Polyporus tenuiculus (=Favolus_ brasiliensis), which
incorporates species with clamped and _ simple-septate
generative hyphae, spores of various sizes, and glabrous to
tomentose pilei. Synonymy has followed the previous work of
Ryvarden on type studies in polypores, together with other
authors' works. I have found several discrepancies regarding
Spegazzini's species, some of which were accepted by
Ryvarden in previous works, but have been changed without
explanation in the present volume. Some species have not
been treated at all, such as Polyporus puttemansii, an alleged
synonym of P. guianensis. Some species are synonymized in a
cryptic way; they are considered as such under the Remarks
or under the Cultural characteristics comments of a species but
are not distinctly cited in its synonymy (e.g., P. hypomelanus
under P. gayanus, and P. lowei under P. admirabilis).

Typographical errors are almost completely absent.
Nevertheless, "mm" has been used instead of ""~m" throughout
the work. [This and other errors are corrected in an errata
sheet: | DiS, Ha] In addition, Spanish characters are omitted.
This is especially unfortunate because it affects the senior
author's name, which is presented as "Nunez" instead of
"Nufiez.". This will almost certainly lead to incorrect
referencing of this work in the literature.

Overall, this monograph sets the basis for a
comprehensive knowledge of the treated genera all over the
world. Discrepancies, such as the cases of recently described
species that are synonymized in this book, will prompt students
to clarify the taxonomic status in several complexes with tools
other than strict morphological criteria. I congratulate the
authors of this work and, recommend it to specialists. Mario
Rajchenberg, Centro de Investigacién y Extensién Forestal
Andino Patagonico, Esquel, Chubut, Argentina.

MYCOTAXON

Volume LX, p. 497 October-December 1996

NOTICE: Transfer of
Professor E. J. H. Corner's Mycological Material

Professor E. J. H. Corner's mycological material, except for the types already in
the holdings of the Cambridge University Herbarium, has been transferred to the Royal
Botanic Garden, Edinburgh (E). It will be housed there along with Professor Comer's
paintings, manuscripts and annotated publications; a small area of the new library extension
has been dedicated to this unique collection of archival material.

As it will take some time to sort and catalogue the material it is impossible to send
specimens out on loan. However anyone wishing to work through collections will be
welcome in Edinburgh where either Evelyn Turnbull or Dr. Roy Watling, Royal Botanic
Garden, Edinburgh EH3 5LR, can guide them through the maze of boxes and containers.

498

Author Index, Volume Sixty

Abdullah, Samir K., J. Guarro and M. J. Figueras New and Interesting
Helicoon Species from Spain. 449-454

Abraham, T. K., K. B. Vrinda and C. K. Pradeep A New Species of
Hygrocybe from India. 361-364

Ammirati, Joseph F., Jr. see Burdsall, Volk and Ammirati

Anderson, Freda, and Virginia Bianchinotti A New Species of Bartalinia Tassi
(Deuteromycotina, Coelomycetes). 191-200

Arambarri, A. M. see Godeas and Arambarri

Baka, Zakaria A., and Halvor B. Gjzrum Egyptian Uredinales. I. Rusts on Wild
Plants from the Nile Delta. 291-303

Baral, Hans Otto Hymenoscyphus seminis-alni, a New Species of the H. fructigenus -
Complex. 249-256

Barr, Margaret E. Planistromelaceae, a New Family in the Dothideales. 433-442

Baydar, Saffet see Sesli and Baydar

Bessette, Alan E. see Fatto and Bessette

Bianchinotti, Virginia see Anderson and Bianchinotti

Boisselier-Dubayle, M. C. see Perreau-Bertrand, Boisselier-Dubayle and
Lambourdiere

Bolontrade, Marcela F. see Wright, Bolontrade and Fazio

Burdsall, Harold H., Jr., Thomas J. Volk and Joseph F. Ammirati, Jr.
Bridgeoporus, a New Genus to Accommodate Oxyporus nobilissimus
(Basidiomycotina, Polyporaceae). 387-395

Buyck, B. see Robert and Buyck

Cano, J. see Castafieda Ruiz, Guarro and Cano

Castafieda Ruiz, Rafael F., J. Guarro and J. Cano Notes on Conidial Fungi. X.
A New Species of Ceratosporella and Some New Combinations. 275-281

, D. E. Fabré, M. P. Parra, M. Pérez and J. Guarro Some Airbome
Conidial Fungi from Cuba. 283-290

see Peldez and Castafieda Ruiz

Cazares, Efrén see Trappe, Jumpponen and Cazares

Chen, Kuei-Yu, and Zuei-Ching Chen A New Species of Thermoascus witha
Paecilomyces Anamorph and Other Thermophilic Thermoascus Species from Taiwan.
225-240

and Thielavia pingtungia sp. nov. a Thermophilic Ascomycete from
Taiwan. 241-247

Chen, Zuei-Ching see Chen and Chen

Claessen De Miranda, Jeanne see Lance Spain and Claessen De Miranda

Currah, R. S. see Tsuneda, Maekawa and Currah

De, A. B. Royoporus —A New Genus for Favolus spathulatus. 143-148

_ see Roy and De

Descals, Enrique see Marvanova and Descals

Diederich, Paul see Etayo and Diederich

Estrada-Torres, A. see Rodriguez-Palma and Estrada-Torres

Etayo, Javier, and Paul Diederich Lichenicolous Fungi from the Western Pyrenees,
France and Spain. IH]. More Deuteromycetes. 415-428

Fabré, D. E. see Castafieda Ruiz, Fabré, Parra, Pérez and Guarro

Fatto, Raymond M., and Alan E. Bessette A New Species of Callistosporium. .
125-128

Fazio, Alejandra T. see Wright, Bolontrade and Fazio

Figueras, Maria José see Abdullah, Guarro and Figueras

see Mercado Sierra, Figueras, Mena Portales
Gjzrum, Halvor B. see Baka and Gjerum

499

Godeas, A. M., and A. M. Arambarri Helicoon septatissimum sp. nov., a New
Species from Tierra del Fuego. 481-484

Guarro, J. see Abdullah, Guarro and Figueras

see Castafieda Ruiz, Guarro and Cano

see Castafieda Ruiz, Fabré, Parra, Pérez and Guarro

Harvey, Alan E. see Larsen, Harvey, Powers and Jurgensen

Hibbett, D. S. Book Reviews 491-496

Hjortstam, Kurt, and Leif Ryvarden New and Interesting Wood-Inhabiting Fungi
(Basidiomycotina—A phyllophorales) from Ethiopia. 181-190

Jumpponen, Ari M. see Trappe, Jumpponen and Cazares

Jurgensen, Martin F. see Larsen, Harvey, Powers and Jurgensen

Koch, J., and K. R. L. Petersen A Check List of Higher Marine Fungi on Wood
from Danish Coasts. 397-414

Lance Spain, Joyce, and Jeanne Claessen De Miranda Scutellospora
cerradensis: An Ornamented Species in the Gagasporaceae (Glomales) from the
Cerrado Region of Brazil. 129-136

and Jeanne Claessen De Miranda Glomus brasilianum: An Ornamented

Species in the Glomaceae. 137-142

Lambourdiere, J. see Perreau-Bertrand, Boisselier-Dubayle and Lambourdiere

Larsen, Michael J., Alan E. Harvey, Robert Powers and Martin F.
Jurgensen A New Species of Botryobasidium (Corticiaceae, Aphyllophorales)
Inhabiting the Interiors of Brown Rotted Logs of Ponderosa Pine. 429-432

Longcore, Joyce E. Chytridiomycete Taxonomy Since 1960. 149-174

Lunghini, D., F. Pinzari and L. Zucconi Studies on Mediterranean
Hyphomycetes. III. Quadracaeamediterranea anam.-gen. and sp. nov. 103-110

Maekawa, N._ see Tsuneda, Maekawa and Currah

Malcolm, W. M. see McCarthy and Malcolm

Manandhar, J. B., and T. W. Mew Pinatubooryzae gen. et sp. nov. and its
Identity During Routine Tests of Rice Seed. 201-212

Marvanova, Ludmila, and Enrique Descals Hyphomycetes from Streams: New
Taxa and New Combination. 455-469

McCarthy, P. M., and W. M. Malcolm Strigulaocculta, a New Saxicolous Lichen
from New Zealand. 323-326

Mena Portales, Julio see Mercado Sierra, Figueras, Mena Portales

Mercado Sierra, Angel, Maria José Figueras, Julio Mena Portales A New
Species of Alysidiopsis from Mexico. 443-448

Mew, T. W. see Manandhar and Mew

Mohammed, C. see Yuan, Mohammed and Wardlaw

Parra, M. P. see Castafieda Ruiz, Fabré, Parra, Pérez and Guarro

Parmalee, J. A. A Catalogue of Types (Uredinales) in the National Mycological
Herbarium (DAOM) Ottawa, Canada. 1-77

Pelaez, Fernando, and Rafael F. Castaiieda Ruiz Amoenodochium humicola
anam. gen. et sp. nov., a New Sporodochial Hyphomycete from Indian Soil. 257-262

Pérez, M. see Castafieda Ruiz, Fabré, Parra, Pérez and Guarro

Perreau-Bertrand, J., M. C. Boisselier-Dubayle and J. Lambourdiere
Mycenasororia sp. nov., Close to M. rosea Gramberg (Basidiomycotina). 263-273

Petersen, K. R. L. see Koch and Petersen

Piccolo Grandi, Rosely Ana, and Derlene Silva Attili Hyphomycetes on
Alchorneatriplinervia (Spreng.)Miill. Arg. Leaf Litter from the Ecological Reserve
Juréia-Itatins, State of Sado Paulo, Brazil. 373-386

Pinzari, F. see Lunghini, Pinzari and Zucconi

Powers, Robert see Larsen, Harvey, Powers and Jurgensen

Pradeep, C. K. see Abraham, Vrinda and Pradeep

Robert, V., and B. Buyck ALLRUS: A System for Standard Description,
Identification and Classification of Russulaceae. 471-480

Roberts, Peter Heterobasidiomycetes from Majorca & Cabrera (Balearic Islands). 111-
123

500

Rodriguez-Palma, M., and A. Estrada-Torres Some Stemonitales (Myxomycetes)
from the State of Tlaxcala, Mexico. 79-102

Roy, Anjali, and A. B. De Taxonomy of Fomitopsis rubidus comb. nov. 317-321

Ryvarden, Leif see Hjortstam and Ryvarden

Sesli, Ertugrul, and Saffet Baydar A Preliminary Checklist of eek in
Turkey. 213-224

Silva Attili, Derlene see Piccolo Grandi and Silva Attili

Trappe, James E., Ari M. Jumpponen and Efrén Cazares NATS Truffle and
Truffle-Like Fungi 5: Tuber lyonii (=T. texense), with a Key to the Spiny-Spored
Tuber Species Groups. 365-372

Tsuneda, A., N. Maekawa and R. S. Currah A New Species of Leptodontidium
from Quercus acutissima.. 485-490

Van der Gucht, Katleen Xylaria Species from Papua New Guinea: Cultural and
Anamorphic Studies. 327-360

Volk, Thomas J. see Burdsall, Volk and Ammirati

Vrinda, K. B. see Abraham, Vrinda and Pradeep

Wardlaw, T. J. see Yuan, Mohammed and Wardlaw

Wright, Jorge E., Marcela F. Bolontrade and Alejandra T. Fazio The Genus
Pyrofomes in Argentina and the Curltural Features of P. lateritius (Aphyllophorales).
305-315

Yuan, Z. Q., C. Mohammed and T. J. Wardlaw Coccoidellaexocarpi sp. nov.
on Exocarpos spp. from Australia. 175-180

Zucconi, L. see Lunghini, Pinzari and Zucconi

INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME SIXTY

501

This index includes names of genera, infrageneric taxa, species, and infraspecific taxa.

New names are in boldface, and new names of suprageneric taxa,
which are also included, are in BOLDFACE
The page on which new taxa are proposed is in boldface.

Acarospora
fuscata 422
Acaulospora
rehmii 141
Acrodictys 275
furcata 278
Actinocladium 275, 278
longiramosum 278
Aecidium
ageratinae 5
berberidis 295
borrichiae 5
chuquiraguae 5
cressae 300
hornotinum 5
ilicina 8
ivae 5
leonense 5
ligustricola 6
millae 6
minimum 6
mundula 6
nectandrae 6
papudensis 6
pereskiae 6
plantaginis 294
poecilochromae 6
pulverulentum 6
pratae 6
punicium 6
sanguinolentum 297-298
tenebrosum 6
Agaricus 264
arvensis 217
augustus 217
bernardii 217
bisporus 217
bitorquis 217
campestris 217
comtulus 217
cupreobrunneus 217
lanipes 217
meleagris 217
rodmani 217
semotus 217
silvicola 217
variecolar 217
xanthodermus 217
xerampelinus 217
Agrocybe
aegerita 217
dura 217
pediades 218
semiorbicularis 217

Allochytridium 151
expandens 151, 160
luteum 151

Allodus
erigeniae 22
megalospora 43

Allomyces 151
attomyces 151
catenoides 151
reticulatus 151

Alternaria
alternata 203

Alysidiopsis 443-445, 447
foliicola 444, 447

lignicola 443, 444-447

pipsissewae 444, 447
yunnanensis 444, 447
Alysidium 447
Amanita 221
caesarea 217
citrina 217
codinae 217
excelsa 217
gemmata 217
junquillea 217
muscaria 217
ovoidea 217
pantherina 217
phalloides 217
rubescens 217
Spissa 217
vaginata 217
var. alba 217
valens 217
verna 217
virosa 217

Amoenodochium 258, 260
humicola 257, 258-260, 262

Amphicypellus 151, 155
elegans 151, 155
Ampullifera
foliicola 416
Amylocarpus

encephaloides 398, 402

Anaeromyces 151, 172
elegans 151, 172
mucronatus 151

Anguillospora 455-456

fustiformis 455-456, 458

Aniptodera 400
Anisostagma
rotundatum 398, 402
Antrodia
juniperina 188

APITALS.

502

Aphanomyces 169
Appendichordella

amicta 398, 402
Arachnophora 275, 278

pulneyensis 278
Arenariomyces

trifurcatus 398, 402
Armillaria 264

bulbosa 218

mellea 218

tabescens 218
Arnaudovia 151

hyponeustonica 151, 167
Arthrobotrys 201-202, 208-209

amerospora 202

anomala 201

constringens 202

dendroides 202

ferox 202

pauca 202

stilbacea 201-202, 209

superba 201
Arthuria

catenulata 7
Arthuriomyces

peckiana 7
Arxiella

terrestris 288
Asperisporium 260, 285

caricae 260, 262
Asteromyces

cruciatus 398, 409
Asterophlyctis

irregularis 151, 172

sarcoptoides 151, 158
Athelopsis

glaucina 182

lembospora 185

Bachmanniomyces
uncialicola 416
Baeodromus
californicus 7
eupatorii 7
holwayi 7
Baeospora
myriadophylla 127
Bartalinia 191-192, 197, 199
bischofiae 197, 199
cordylines 199
lateripes 199
mellea 191-193, 194, 196-199
pistacina 199
robillardoides 197-198
tamarindi 197
Basidiodendron
cinereum 116
eyrei 117
nodosum 111, 114-117
rimulentum 111, 114, 116-117
Beauveria 209
tenella 209

Biconiosporella
corniculata 398-399, 402
Biflua
physasca 398, 403
Blastocladia 169-171
arborata 151
caduca 151
coronata 151
cristata 151
didyma 151
elegans 151
excelsa 151
filamentosa 152
fruticosa 152
fusiformis 152
globosa
var. minutissima 152
heterosporangia 152
mammilata 152
picaria 152
pileota 152
pusilla 152
sessilis 152
spiciformis 152
Blastocladiella 152
Subg. Blastocladiella 152
Subg. Cystocladiella 152, 157
Subg. Eucladiella 152, 159
anabaenae 152
britannica 152
colombiensis 152
nova-zeylandiae 152
Blastocladiopsis
elegans 152
Blyttiomyces
aureus 152
conicus 152
exuviae 152 -
gregarum 152
harderi 152
laevis 152
lenis 152
rhizophlyctidis 152
spinosus 152, 169
vaucheriae 153, 166
Bolbitius
vitellinus 218
Boletus
aereus 215
aestivalis 215
calopus 215
chrysenteron 215
edulis 215
elegans 215
erythropus 215
luridus 215
porphyrosporus 215
queletii 215
radicans 215
regius 215
retipes 215
rufus 215

[Boletus] satanoides 215

versicolor 215

zelleri 215
Botryobasidium 429-431

curtisii 431

isabellinum 120

medium 429, 431

ponderosum 429, 430-431

subcoronatum 429, 431
Botryorhiza

hippocrateae 7
Botryotrichum

peruvianum 288
Bovicornua

intricata 398, 403
Bridgeoporus 387-388, 390

nobilissimus 390, 393-394
Bullaria

carthami 293
Byssoloma 416

leucoblepharum 416

subdiscordans 416

Caecomyces 153
communis 153
equi 153

Callimastix
cyclopis 153
frontalis 153, 161

Calliospora
diphysae 77
holwayi 77

Callistosporium 125
luteo-olivaceum 125, 127
purpureomarginatum 125-127
rubescentifolia 125

Calocera
cornea 122

Camarophyllus
subradiatus 216
virgineus 216

Canocybe
coprophila 218
lactea 218
rickeniana 218

Canteria 153
apophysata 153, 164

Capronia
ciliomaris 398-399, 403, 414

Carbosphaerella
leptospharioides 398, 403
pleosporoides 398, 403

Catenaria
auxiliaris 153
indica 153
ramosa 153
spinosa 153
uncinata 153
vermicola 153
verrucosa 153

Catenochytridium
carolineanum

f. marinum 153
hemicysti 153
marinum 153
oahuense 153
Catenophlyctis 153

peltata 153

variabilis 153, 159, 163, 166

var. olduvaiensis 153

Caulochytrium 153

gloeosporii 153

protostelioides 154
Ceraceopsora

elaegni 8
Ceratium

hirundinella 170
Ceratobasidium 117
Ceratosebacina

longispora 111, 117-118
Ceratosporella 275, 278-279

basibicellularia 278-279

basicontinua 278-279

bicornis 278-279

caliculata 278-279

cheiroidea 278-279

compacta 275, 276-280

deviata 278

disticha 278-279

fertilis 278-279

flagellifera 278-279

goidanichii 278

lambdaseptata 278

longiramosa 278

novae-zelandiae 278-279

ponapensis 278-279

pulneyensis 278

stipitata 278-279
Ceriosporopsis

caduca 398-400, 403

cambrensis 399

halima 398, 403

hamata 400

longissima 399

tubulifera 398, 403
Cerotelium

holwayi 8
Chaetomastia

typhicola 398-399, 404
Chaetopsina

ramifera 379
Chaetosphaeria

chaetosa 398, 404
Chalara

aurea 373, 375-376
Chloridium

virescens 373

var. virescens 373, 375-376

Chroogomphus

rutilus 215
Chrysocyclus

mikaniae 8
Chrysomyxa

ilicina 8

503

504

[Chrysomyxa] ledi
var. glandulosi 8
var. groenlandici 8
var. vaccini 9
monesis 9
weirll 9
Chrysopsora
gynoxidis 9
mikaniae 8
Chytridium 154, 157, 169
adpressum 154
aggregatum 154, 157
apophysatum 154
brevipes 154, 157
cejpii 154, 157
chlorobotryis 154, 157
citriforme 154, 157
closterii 154
codicola 154, 157
coleochaetes 154, 169
confervae 154-155
corniculatum 154
cresentum 154
deltanum 154
fusiforme 154
gibbosum 154-155, 157
hemicysta 153
horariumforme 154
hyperparasiticum 154
inflatum 154, 157
isthmiophilum 154, 157
kolianum 154, 158
lagenaria 154, 158
var. japonense 158
latipodium 154
mallomonadis 155, 158
marylandicum 155
mucronatum 155, 158
neopapillatum 155
oedogonii 155, 158
olla 154
ottariense 155
parasiticum 155-156
pilosum 155
proliferum 155
reniforme 155
schenkii 154-155, 158
var. dumontii 155
scherffelii 155, 158
sexuale 155, 158
sparrowii 155
stellatum 155, 158
suburceolatum 156
telmatoskenae 155
turbinatum 155, 158
Chytriomyces 151, 155
angularis 155
annulatus 155
confervae 154-155
cosmarii 155
elegans 151, 155
gilgaiensis 155

heliozoicola 156
hyalinus 156

var. granulatus 156
laevis 156
macro-operculatus 156

var. hirsutus 156
mammilifer 156
mortierellae 156
multi-operculatus 156
nagatoroensis 156
poculatus 156
reticulatus 156
reticulosporus 156
rhizidiomycetis 156
rotoruaensis 156
suburceolatus 156
vallesiacus 156
verrucosus 156
willoughbyi 155-156

Ciboria
alni 254
seminicola 254
uliginosa 254
Cirrenalia

macrocephala 398-399, 409

Cladochytrium 162
aureum 156
granulatum 156, 162
indicum 156
novoguineense 156
replicatum 156
salsuginosum 156

Cladonia 416
furcata 416
parasitica 415, 424-425
stellaris 423
uncialis 416

Cladosporium 154, 377, 447

cladosporioides 373, 376-377

Clitocybe
alexandri 218
cyathiformis 218
dealbata 218
ectypa 218
familia 218
fragrans 218
geotropa 218
gibba 218
gigantea 218
infundibuliformis 218
inversa 218
laccata 218
lignatilis 218
nebularis 218
odora 218
paropsis 218
perilis 218
radicellata 218
rhizophora 218
squamulosa 218
tabascens 218

Clitopilus
prunulus 216
Coccodothella 175
Coccoidea
tsurudae 438
Coccoidella 175-176, 178-179, 438
exocarpi 175-176, 179
euglypta 179
fici 179
fusispora 179
kawagoiil 438
perseae 178-179
pulvinata 179
quercifolia 179
scutula 175, 178-179
spinulosae 438
stevensonia 179
tsurudae 438
Codinaea
simplex 379
Coelomomyces 150
Coelomycidium 156
Colacogloea
peniophorae 114
Coleosporium
adenocaulonis 9
anceps 9
begoniae 9
cheoanum 10
choerospondiatis 10
crowellii 10
elephantopodis 10
paraphasatum 10
rubiicola 10
steviae 10
verbesinae 10
viguierae 10
Collaria 98, 100
rubens 98
Colloderma 100
roblustum 82
Collybia
butyracea 218
cessans 218
colera
var. rubescentifolia 125
dryophila 218
fusco-purpurea 219
grammocephala 219
longipes 219
peronata 219
tuberosa 219
Columnodontia 184
Comatricha 100
aggregata 97
elegans 97
laxa 96
lurida 96-97
nigra 97
rubens 79, 98
subcaespitosa 98-100
tenerrima 100

505

Coniothyrium
ramalinae 418
Coprinus
atramentarius 216
comatus 216
disseminatus 216
galericuliformis 216
micaceus 216
niveus 216
ovatus 216
Ppicaceus 216
radiatus 216
Cornutispora
lichenicola 418
Corollospora
lacera 398-399, 404
luteola 398-399, 404
maritima 398, 404
Cortinarius
auroturbinatus 220
bulbosus 220
bulliardii 220
cotoneus 220
glaucopus 220
malachioides 220
olivaceo-fuscus 220
punctatus 220
purpurascens , 220
subturbinatus 220
subvalidus 220
Crepidotus
cesatii 220
mollis 220
pubescens 220
sphaerosporus 220
Crinigera
maritima 399
Cronartium
occidentale 10
ribicola 11
Cryptophiale
kakombensis 373, 376-377
udagawae 373, 377-378
Cudoniella
clavus 254
Cumminsiella
wootoniana 11
Cumulospora
marina 398-399, 409
Curvularia
tuberculata 288-289
verrucosa 288

Cylindrochytrium
endobioticum 157
Cylindrocladium
scoparium 373, 377-378
Cystoderma

amianthinum 217

Dacrymyces 112
enatus 122
stillatus 122

506

Dactylaria
kumamotoensis 288-289
Dactylomyces
crustaceus 230
Dangeardia 157
appendiculata 157, 172
echinulata 157
molesta 157, 159
ovata 157
sporapiculata 157
var. minor 157
Dangeardiana 157
apiculata 157-158
eudorinae 157
leptorrhiza 157, 172
sporapiculata 157
Dextrinodontia
molliuscula 182
Diacheopsis 83, 100-101
effusa 83
insessa 79, 83-84, 88
nannengae 83
Diaporthe 435
Diatrype
acervata 434
Dicheirinia
superba 11
Dictyochaeta
simplex 373, 379, 382
Dictyomorpha 157
dioica 157, 167
var. pythiensis 157
Dictyosporium
pelagicum 398-399, 409
toruloides 398-399, 410
Didymella
myricariae 439
Didymopsora
chuquiraguae 11
triumfettae 11
Dietelia
eupatoril 7
holwayi 11
Digitatispora
marina 398, 411
Diplochytridium 154, 157
aggregatum 154, 157
brevipes 154, 157
cejpit 154,157
chlorobotryis 154, 157
citriforme 154, 157
codicola 154, 157
gibbosum 154, 157
inflatum 154, 157
isthmiophilum 154, 157
kolianum 154, 158
lagenarium 154, 158
var. japonense 158
mallomonadis 155, 158
mucronatum 155, 158
oedogonii 155, 158
schenkii 155, 158

scherffelii 155, 158
sexuale 155, 158
stellatum 155, 158
turbinatum 155, 158
Diplodia
oraemaris 398, 410
Diplodina
spinulosae 438
Diplomitoporus
rimosus 181, 188
Diplophlyctis 158
asteroidea 158
butermerense 162
buttermerensis 158, 162
chitinophila 158
complicata 158, 162
intestina 172
nephrochytrioides 158
sarcoptoides 151, 158
sexualis 158, 162
versiformis 158
Dothidella 435
Dothistroma 437
septospora 437
Dryosphaera
navigans 398-399, 404
Dwalomyces
taiwanensis 288

Edythea
tenella 12
Eichleriella
deglubens 123
Eiona
tunicata 398, 404
Endochytrium
cystarum 158
multiguttulatum 158
Endophragmiella 103
boewei 106-107
verticillata 107
Endophyllum
alaskanum 12
holwayi 11
lacus-regis 12
Endothia
parryii 435
Enerthenema 100
papillatum 83
Entoloma
clypeatum 216
lividum 216
Entophlyctis 158
apiculata 157-158
bulligera 158-159
var. brevis 159
caudiformis 159

confervae-glomeratae 158-159

crenata 159
filamentosa 159
helioformis 159
lobata 159

[Entophlyctis] luteolus 159
mammilliformis 159
molesta 157, 159
obscura 159
reticulospora 159
rhizina 159
sphaerioides 159
texana 159
variabilis 153, 159, 166-167
vaucheriae 159
willoughbyi 159

Eriomycopsis 209-210
minuta 209

Eruptio 433, 437, 440
acicola 433, 438
pini 433, 438

Eupenicillium 476

Exidiopsis
effusa 123

Favolus 145, 147
brasiliensis 145, 147
moluccensis 145, 147
spathulatus 143, 145, 147
tenuiculus 145, 147

Flammula
hybrida 219
liquiritiae 219
penetrans 219
sapinea 219
spumosa 219
velutipes 219

Flavoparmelia
caperata 422-423, 426-427

Fomes 388-389
fulvo-umbrinus 306, 314
juniperinus 306
lateritius 305, 307
nobilissimus 390
perlevis 311

Fomitopsis 320
pinicola 320
rubidus 317, 319

Fusarium 209-210, 417
peltigerae 416

Fusicladium 285

Gaertneriomyces 159
semiglobiferus 159, 165-166
tenuis 159

Galerina
marginata 220
unicolor 220

Gerlachia
oryzae 209

Gloeosporium 153

Gloiothele 182
lamellosa 182

Glomus 137, 140
albidum 141
brasilianum 137-138, 139-141
callosum 141
clarum 141

convolutum 137
leptotichum 141
maculosum 141
manihotis 137
occultum 137
pallidum 141
scintillans 141
Gonapodya
polymorpha 170-171
Gonopodya 159
terrestris 159
Groenhiella
bivestia 398, 404
Gymnoconia 7
peckiana 7
Gymnosporangium
asiaticum 13
bermudianum 12
betheli 12
connersii 12
davisii 12
durum 13
ellisii 13
fraternum 13
gaeumanii 13
ssp. albertensis 13
haraeanum 13
harknessianium 13
nelsonii 13
nootkatense 13
Gyoerffyella 463

Hainesia 418
lythri 418
pertusariae 415, 417
Haligena
elaterophora 398, 405
salina 398-399, 405
Halosarpheia 400
retorquens 400
trullifera 398, 405
viscosa 398, 400, 405
Halosphaeria
appendiculata 398-400, 405
Halosphaeriopsis
mediosetigera 398, 405
Hamaspora
longissima 14
ozeenis 14
Hamatocanthoscypha 255
Hapalopera 159
achnanthis 160, 168
difficilis 160, 169
fragilariae 160, 170
melosirae 160, 170
piriformis 160, 170
Haplographium
heliocephalum 379
Haplotrichum 429-431
curtisii 431
Harpochytrium 150, 160
adpressum 160

507

508

[Harpochytrium] apiculatum 160
botryococci 160
hedenii 160
hyalothecae 160
intermedium 160
monae 160
natrophilum 160
ornithocephalum 160
tenuissimum 160
viride 160

Hebeloma
leucosarx 220
populinum 220
sinapizans 220

Helicodendron
japonicum 451

Helicogloea
lagerheimii 123

Helicoon 449, 451, 481, 483-484
auratum 484
chlamydosporum 451, 454, 484
ellipticum 484
farinosum 451, 484
fuscosporum 449, 451, 453-454, 484
macrosporum 451
microsporum 449, 450-452
peruamazonense 451
pluriseptatum 484
richonis 483-484
septatissimum 481-484
sessile 451, 484

Helotium
nyssicola 254-255

Hemileia
oxyanthi 14

Heterobasidion
annosum *430

Hobsonia
christiansenii 421-422

Hohenbuehelia
atrocoerulea 219
rickenii 219

Hyalopsora
adianti-capilli-vernis 14
diplazii 14
polypodii 14

Hyaloscypha 255

Hygrocybe 361, 364

Sect. Coccineae 364
conica 216
langei 216
nigrescens 216
parvispora 361, 363
subglobispora 216

Hy grophorus
calophyllus 216
chrysodon 216
croceus 217
eburneus 217
fuscoalbus 217
pudorinus 217
russula 217

Hymenoscyphus 249-250, 255
calyculus 255
caudatus 255
fructigenus 249, 253-254, 256
var. aesculicarpa 254
var. carpini 253-254
var. coryli 254
imberbis 455-456
malawiensis 456, 458
rokebyensis 254
scutula 255
seminis-alni 249-250, 253-256
Hyphoderma
argillaceum 113-114
praetermissum 114
Hyphodontia
spathulata 182
Hypholoma
fasciculare 218
Hypocrea
agaves 435
Hypogymnia
subphysodes
var. austerodioides 175
Hypotrachyna
revoluta 422-423, 426-427

Idriella
falcata 373, 378-379, 383
Illosporium 422
aurantiacum 421
corallinum 420, 422
roseum 421
Ingoldia 463
Inocybe
fastigiata 220
fibrosa
var. trivialis 220
geophylla
var. lilacina 220
gymnocarpa 220
patouillardii 220
perbrevis 220
subrubens 220
xanthodermus 221
Itersonilia
perplexante 172

Karlingia 151, 160
aurantiaca 160
exo-operculata 160
expandens 151, 160
granulata 160
lacustris 160
lobata

var. microspora 161
polonica 161
rosea 156, 169
spinosa 161

Karlingiomyces 160-161
granulatus 160
laevis 161

Kavinia

alboviridis 183
Kellermania

anomala 434

uniseptata 434

yuccifoliorum 434

yuccigena 434
Kionochaeta

ramifera 373, 378-379, 383
Kirschsteiniothelia

maritima 398, 405
Kochiomyces 161

dichotomus 161, 165
Kohlmeyeriella

tubulata 398-399, 405
Kuehneola

ramacharii 15
Kuehneromyces

mutabilis 218

Laccaria
amethystina 219
laccata 219
Lacrymaria
lacrymabunda 216
var. velutina 216
Lactarius 471
Lacustromyces 161
hiemalis 161
Lamproderma 100
columbinum 95
echinulatum 95
nigrisplendidum 95
scintillans 96

Lasallia

papulosa 423
Laschia

spathulata 143, 145, 147
Lauriomyces

heliocephala 373, 379-380, 383-384

Lautisporopsis
circumvestita 398, 406, 414
Lecanosticta 437
acicola 437
Leccinum
carpini 215
corsicum 215
insigne 215
oxydabile 215
quercinum 215
scabrum 215
subleucophaeum 215
versipelle 215
Lentescospora
submarina 400
Lentinula
edodes 486, 488
Lepiota
acutesquamasa 217
alba 217
badhamii 217
brunneo-incarnata 217

cristata 217

cygnea 217

holosericea 217

jubilaei 217

litoralis 217

naucina 217

rhodorrhiza 217
Lepista :

luscina 219

nuda 219

saeva 219

sordida 219
Leptodontidium 485-486

elatius 485

irregulare 488

orchidicola 485-486, 488, 490

quercuum 485-486, 488, 490
Leptogium 325
Leptosphaeria

oraemaris 398, 406

pelagica 398-399, 406
Leucoagaricus

leucothites 217
Leucopaxillus

amarus 219

cerealis 219

gentianeus 219
Lichenoconium

cargillianum 418
Lichenopuccinia

poeltii 418
Limacospora

sundica 398, 406
Lindra

inflata 398, 406
Litschauerella

abietis 183

clematitis 183

gladiola 183
Loratospora 433, 437, 440

aestuarii 437
Loweporus 306, 314
Lulworthia

fucicola 398-399, 406

lignoarenaria 398, 406

Macrochytrium
botrydiella 161
botrydioides

var. minutum 161

Macrolepiota
excoriata 217
mastoidea 217
permixta 217
procera 217

Macrophomina
phaseolina 419
pseudeverniae 415, 419-420

Mainsia
clara 15
holwayii 15
tenella 15

» 509

510

Marasmius

androsaceus 219

calopus 219

caryophylleus 219

exocarpi 175

oreades 219

perforans 219

ramealis 219

resinosus 219

rotula 219

splachnoides 219
Marchandiomyces 415, 420, 423, 427

aurantiacus 415, 421, 423

corallinus 415, 420-423
Marinosphaera

mangrovei 398-400, 406, 413-414
Marinospora

calyptrata 398-399, 407

longissima 398-400, 407
Marisolaris

ansata 398, 407
Massarina

thalassiae 398-400, 407, 414
Melampsora 291-292

aleuritidis 15

epitea

f. sp. tsugae 15

euphorbiae 291-292

euphorbiae-gerardiana 291

helioscopiae 291

lini 15

piscariae 15
Melampsorella

kriegeriana 16
Melanelia 422-423

exasperata 422-423

glabratula 422-423

panniformis 422-423

substygia 427

tominii 427
Melanoleuca

cinerascens 219

cognata 219

Meliola
exocarpi 175
Melzericium
udicola 183
Menegazzia
confusa 175
Metacapnodium

juniperi 439
Microallomyces 161
dendroideus 161

Microcyclus 175-176, 178, 433, 438, 440

angolensis 438

kawagoelii 433, 436, 438

porlieriae 438

tinctorius 178, 438
Microdochium

falcatum 379
Micromphale

brassicolens 219

Micromyces
furcata 161
grandis 161
Micronegeria
alba 16
Milesia
asplenii-incisa 16
vogesiaca 17
Milesina
asplenii-incisa 16
blechnicola 16
faulliana 16
feurichii 16
jezoensis 16
kameiana 16
kriegeriana 16
laeviuscula 17
tobinagai 17
vogesiaca 17
Milospium 425
graphideorum 425
lacoizquetae 415, 424-425
Mimema
holwayii 66
Mitochytridium
regale 161
Monoblepharella 161
Monoblepharis
micrandra 161
Monodictys
pelagica 398-399, 410
Monophagus 161
blackmanii 161
bruhlii 161
Morchella 263
Mortierella 156
Muellerella 439
dalbergiae 439
Mycena 263-268
Sect. Calodontes 263-264, 267
Subsect. Purae 264, 268, 271
alcalina 219
atroalba 219
auroricolor 271
corticola 219
epipterygia 219
fibula 219
filopes 219
galericulata 219
inclinata 219
integrella 219
niveipes 219
parabolica 219
plicosa 219
polygramma
f. pumila 219
pseudocorticola 219
pura 219, 263-268, 271
var. rosea 264
f. pura 271
f. roseoviolacea 271
rosea 263-268, 270-271

[Mycena] sanguinolenta 219
sororia 263-268, 269, 271
speirea 219
strobilicola 219
stylobates 219
subincarnata 219
tintinnabulum 219
vulgaris 219

Mycoacia 181, 184
aurea 185
austro-occidentalis 185
brunneofusca 181, 183, 185
columellifera 184
fuscoatra 183-184
lutea 184
nothofagi 184
subceracea 184
subconspersa 185
subfascicularia 185
uda 184

Mycoleptodiscus
brasiliensis 373, 380-381

Mycosisymbrium
cirrhosum 262

Mycosphaerella 435, 437, 440
apophlaeae 440
ascophylli 440
dearnessii 437
deightonii 435
gibsonii 437
myricariae 439
pini 437-438

Mycosphaerellopsis 433, 439-440
myricariae 436, 439

Myxarium
nucleatum 112, 123

Naematoloma
sublateritium 218
Nais
inornata 398, 407
Naucoria
abstrusa 221
Nautosphaeria
cristaminuta 398, 407, 414
Neocallimastix 161
frontalis 153, 161-162
hurleyensis 162
joyonil 162, 164
patriciarum 161-162
variabilis 161-162
Nephrochytrium
bipes 162
buttermerense 158, 162
complicatum 158, 162
sexuale 158, 162
Nereiospora
comata 398-399, 407
cristata 398, 407
Nia
vibrissa 398, 411
Nothoravenelia

japonica 17
Nowakowskiella 162
crassa 155, 162
delica 162
elegans 162
endogena 162
granulata 156, 162
keratinophila 162
methistemichroma 162
moubasherana 162
multispora 162
pitcairnensis 162
profusa 162, 171
f. constricta 162
sculptura 162
Nyssopsora
koelreuteriae 17

Obelidium
megarhizum 162

Occultifur 112

Ocostaspora

apilongissima 398, 400, 408

Oedogoniomyces 163
Oidium 431
curtisil 431
Oligoporus 389
Oligostroma
acicola 437-438
Olpidium 163, 166
appendiculatum 163
bornovanus 163, 167
brassicae 163, 167
cucurbitacearum 163, 167
entophlyctoides 163
fulgens 163
incognitum 163
indicum 163
indum 163
longum 163
nematodae 163
poreferum 163
pseudoeuglenae 163
radicale 163, 167
rostriferum
var. indica 163
sparrowli 163
synchytrii 163
vermicola 163
virulentus 163, 167
wildemani 163, 167
zopfianus 163, 167
Omphalina
abiegna 219
gracillima 219
hepatica 219
hydrogramma 220
leucophylla 220
olearius 220
velutina 220
Omphalotus
olearius 215

511

12

Ondiniella
torquata 398-399, 408, 414
Oogaster 370
Orbimyces
spectabilis 398, 410
Orpinomyces 164
bovis 164
intercalaris 164
joyonii 162, 164
Oudemansiella
mucida 220
radicata 220
Oxyporus 387-389
nobilissimus 387-390, 394
populinus 389

Pachycladina

mutabilis 456
Paecilomyces 225-226, 230, 232

crustaceus 235

taitungiacus 226-227
Panaeolus

acuminatus 216

campanulatus 216

fimicola 216

foenisecii 216

retirugis 216
Panellus

mitis 220

stypticus 220
Pannaria

inmixta 325
Parmelia 415, 417, 419, 422, 426

hygrophila 419

omphalodes 422-423

perforata 418

saxatilis 418-419, 422-423, 426

sulcata 418-419, 422-423, 426
Parmeliella

testacea 416
Parmelina

quercina 422-423
Parmeliopsis

ambigua 422-423
Parmotrema

crinitum 422-423
Passalora 285
Paxillus

atrotomentosus 215

involutus 215

panuoides 215

rhodaxanthus 215
Pazschkeella

brasiliensis 438
Peltigera

rufescens 416
Penicillium 476
Peniophora 187

quercina 187
Penzigia

frustulosa 338
Perenniporia 306, 314

gomezii 306, 314
Peridermium
yamabense 17
Pertusaria 415, 417-418, 427
albescens 427
multipuncta 422-423, 427
ophthalmiza 427
pertusa 422-423, 427
pustulata 422-423
Pestalozzina
cordylines 199
Phaeoisaria
clematidis 373, 380-381, 383
infrafertilis 373, 380-381, 383
Phaeophyscia 421
orbicularis 421
Phakopsora
cheoana 18
ehretiae 18
meliosmae 19
psoraleae 18
tecta 18
Phanerochaete 184-185
rosea 185
Phellinus
demidoffii 306
Phlebia 184
Phlebiella
ardosiaca 185
Phlegmacium
amoenolens 221
variecolar 221
Phlyctidium 164, 168
anatropum 164, 168
apophysatum 153, 164
brevipes
var. marinum 164, 169
bumilleriae 164, 169
globosum 164, 171
keratinophilum 164, 169
var. savulescui 164
marinum 164, 170
mycetophagum 164, 170
olla 164, 170
scenedesmi 164, 171
spinulosum 164, 171
tenue 164, 171
tubulatum 164
Phlyctis 417
Phlyctochytrium 173
acuminatum 164, 173
aestuarii 164, 168
africanum 164, 173
apophysatum 164
arcticum 164, 173
aureliae 165-166
biporosum 169
californicum 159, 165
chandleri 165
circulidentatum 165
cystoferum 165, 167
dichotomum 161, 165

[Phlyctochytrium] dissolutum 165
furcatum 165
hirsutum 165
incrustans 165
indicum 165
irregulare 165
irregularis 165
kniepii 165, 173
lackeyi 165
macrosporum 165
mangrovii 165
marilandicum 165
megastomum 165
mucosum 165
multidentatum 165
neuhausiae 165
palustre 165, 173
parasitans 165
peruvianum 165, 173
planicorne 166, 169
plurigibbosum 166, 173
powhatanense 166
punctatum 166, 173
recurvastomum 166
reinboldtae 166
semiglobiferum 159, 166
spinosum 166
unispinum 167
variable 165-166
vaucheriae 153, 166
verruculosum 166, 173

Phlyctorhiza
peltata 153
variabilis 153, 159, 166

Pholiota
adiposa 218
astragalina 218
aurivella 218
caperate 218
destruens 218
flammans 218
radicosa 218
spectabilis 218
squarrosa 218
tuberculosa 218

Phragmidium
andersoni 18
boreale 19
fusiforme

var. novi-boreale 18
guatemalense 18
itoanum 18
ivesiae

ssp. wyomingense 19
longissima 14

Phragmopyxis
acuminata 19

Physalidiopsis 107

Physcia 421-422
adscendens 421-422
aipolia 422-423
caesia 421-422

a3

semipinnata 421-422
stellaris 421
tenella 421
Physisporinus 389
rivulosus 181, 188
Physoderma 149-150
Physopella
meliosmae 19
Pileolaria
clemensiae 19
pistaciae 19
Pinatubo 201-203, 204, 209
oryzae 201-203, 204-206, 208-210
Piptarthron
crassisporum 437
pluriloculare 435
uniloculare 437
Piromonas
communis 166
Piromyces 166
communis 166
dumbonicus 166
mae 166
minutus 166
rhizinflatus 166
spiralis 166
Planistroma 433, 435, 440
nolinae 437
obtusilunatum 435
yuccigenum 435
Planistromella 433-435, 440
acervata 433-434, 436
parryi 433, 435-436
torsifoliorum 434
uniseptata 434
yuccifoliorum 434
yuccigena 433-434
PLANISTROMELLACEAE 433, 436,
438, 440
Platismatia
glauca 418
Pleotrachelus 163, 166
askaulos 166
bornovanus 163, 167
brassicae 163, 167
fulgens 163
virulentus 163, 167
wildemani 163, 167
zopfianus 163, 167
Pleuropedium 455, 458
multiseptatum 455, 458, 460, 462, 467
tricladioides 458, 460, 463-464, 466-
467
Pleurotus 264
cornucopiae 215
dryinus 215
eryngii 215
ilgazicus 215
nemecii 215
ostreatus 215
var. salignus 216
pubescens 216

514

[Pleurotus] pulmonarius 216
serotinus
var. flaccidus 216
ulmarius 216
Plowrightia 435
agaves 435
circumscissa 435
williamsoniana 435
Plurisperma
dalbergiae 439
Pluteus
atricapillus 216
cervinus 216
minutissimus 216
petasatus 216
salicinus 216
tomentosulus 216
Podochytrium
chitinophilum 167
dentatum 167
ellerbeckense 167
Polypaecilum 230
Polyphagus 151
asymmetricus 167
elegans 167
hyponeustonica 151, 167
serpentinus 167
starrii 167
Polyphlyctis 167
cystofera 165, 167
unispina 167
Polyporus 145, 147, 264, 320
connatus 389
demidoffii 305
moluccensis 145, 147
populinus 389
rubidus 317, 320
spathulatus 147
Polystomella
kawagoli 438
Polythrincium 260
trifolii 260, 262
Porina 325
guentheri 325
leptalea 325
partita 325
Postia 389-390
Powellomyces 158, 167
hirtus 167
variabilis 159, 167
Pringsheimiella 157
dioica 157, 167
Prospodium
amphilophii 19
anomalum 20
arrabidaeae 20
cremastrum 20
cyathiforme 20
depallens 20, 41
holwayi 20
impolitum 20
lundiae 20

palmatum 20
permagnum 20
reticulatum 20
stizophyllii 20
tecomicola 21
transformans 41
Psalliota
edulis
var. valida 217
perrara 217
Psathyrella
candolleana 216
fatua 216
gracilis 216
melanthina 216
obtusata 216
prona 216
spadiceogrisea 216
Pseudevernia
furfuracea 415, 419
Pseudopileum 167
unum 167
Pseudosphaerella 439
Psilocybe
pratensis 218
Puccinia 3, 291
abramsii 29
abrupta 29
var. partheniicola 30
absicca 29
acarnae 25
achilleae 23
actinostemonis 45
additicia 30
aegopogonis 3, 50
aemulans 30, 32
aenigmatica 47
affinis 30
var. triporosa 30
agnita 55
agnitionalis 39
agoseridis 27
albicera 47
albigensis 295
albula 23
albulensis
ssp. cascadensis 59
alia 23
allandabilis 39
alnetorum 297
altissimorum 25
amphigena 50
amphilophii 19
amphispilusa 54
amplifica 47
anguriae 43
anisicanthi 21
antioquiensis 43
arabicola 3, 41
arisanensis 77
aristidae
var. chaetaiae 50

[Puccinia ] asiatica 56
asprellae-japonicae 50
aulica 61
austroberingiana

ssp. austroberingiana 56

ssp. saxifragarum 57
axiniphylii 30
baccharidis-spartea 23
baeumleriana 23
balsamorhizae 30
basiporula 28
batesiana 30
benokiyamensis 54
berberis-trifoliae 40
bessei 49
blutanensis 59
boliviana 30
brachycomes 23
brachytela 31
burnettii 50
butleri 27
caborcensis 31
calanticariae 31
calcitrapae

var. centaureae 293
caleae 31

var. cuernavacae 31
californicae 25
calliquensis 46
calvatica 47
canadensis 62
cancellata 292
Caricis-japonicae 43
caricis-shepherdiae 43

’ carthami 293

cenchri

var. africana 293
centaureae 26

var. diffusa 26
cera 48
chelonis 59
chisosensis 50
chloracae 31
cinnamomicola 48
claytoniicola 55
cleomis 42
clintonii

var. albae 59

var. bracteosa 59

var. clintonii 59

var. groenlandicae 59

var. ornithorhyncae 59

var. racemosa 59

var. sylvaticae 60
coccocypseli 56
codyi 41
cognata

var. cognata 31

var. echinulata 31

var. fraseri 31
compositarum 25
concumulata 56

od

confraga 60
congdonii 43
conjuncta 62
connersii 51
consueta 24
consulta 24
conturbata 47
coronospora 48
corylopsidis 46
cressae 300
cruciferarum

ssp. borealis 41

ssp. nearctica 42

ssp. wyomingensis 42
cryptandri

var. luxurians 50
cupheae 49
cyani 26

var. sublevis 26
cymbopogonis 50
cymopteri 22

ssp. oregonensis 22
cynodontis 291, 293-294
cyperi 44
cyperi-tegetiformis 291, 293
cyrnaea 292
dasypyri 298
decpecanea 39
defecta 38
depallens 20, 41
desantherae 60
desmanthodii 32
detonsa 42
dianthi-japonica 42
dichelostemmae 21
diplachnicola 51
discors 22
discreta 39
dispersa 298
distenta 46
distinguenda 42
distorta 47
douglasii

var. douglasii 53

var. hoodii 53
drabicola 42
effusa 62
egmontensis 38
elaeagni 45
electrae 30

var. depressiporosa 32

var. electrae 32

var. robusta 32
ellisiana SO
enceliae 30

var. aemulans 32-33

var. enceliae 32, 37
erigeniae 22
eriophorii

var. apargidii 44
eriophyllii 29
eriostemonis 56

516

[Puccinia ] espinosarum 28 var. meridionalis 57
excursionis 24 var. minor 57
eximia 56 var. saxifragae-micranthae 57
exitiosa 41 hieracii 27
expetiva 24 holwayula 33
fausta 39 huteri 58
ferox 32 hyalina 44
festata 45 idonea 39
fidulis 47 illatabilis 39
fimbristylidis 44 impedita 47
fischeri 57 imperatae 291, 295
fragilis impetrabilis 39

var. hoodii 53 impolita 24

var. fragilis 53 improcera 24
fragosoana 291, 293, 295 improvisia 40
flavo-virens 44 inaequata 40
franseriae 32 inanipes 28
fraseri 27 inaudita 33
fumosa 54 inclusa
fundata 39 var. boreohesperia 26
fuscata 47 var. brevifolia 26
gahniae 44 var. flodmannii 26
ghiesbrechtii 33 var. mexicana 26
giliae 54 incomposita 24
giliicola 54 indagata 24
gilliesiana 33 inopina 24
gilva 41 inrecta 49
glacieri 62 insignis 43
globulifera 33 insperata 27
glumarum 300 interjecta 24
grahami 22 intermixta 34
graminis invelata 34

ssp. graminicola 295 iostephanes 34

ssp. graminis 295 irregularis 34
grenfelliana 22 irrequisita 26
grumosa 49 isiacae 291, 296
guardiolae 33 juncellii 44
guatemalenses 33 karelica
gulosa 54 ssp. laurentia 44
gymnandrae kuhniae

ssp. alascensis 60 var. robusta 28

ssp. camschatcensis 60 kukkonensis 49

ssp. gymnandrae 60 kuntzii 40

ssp. yunnansis 60 kwangsiana 26
gymnolomiae 32-33 kweichowana 55
gypsophila-repentis 42 lagenophorae 291, 296
haleniae 45 laschii
harae 57 var. fennica 26
harknessii var. palustris 26

var. harknessi 27 var. pannonici 27

var. stephanomeriae 27 var. xerophila 27
hedysari-paniculati 72 lauricola 48
helianthellae 33 lepturi 50
helianthi 33 lezpedezae-procumbentis 72
helicalis 60 lisianthi 46
heterothalami 24 loranthicola 49
heucherae machii 48

var. asiatica 56 magnusiana 297

var. austroberingiana 56 mahoniae 40

var. cordillerana 57 majuscula 38

var. diffusistriata 57 mandevilleae 23

var. japonica 57 mariae 62

[Puccinia ] marie-wilsonii

var. montiae 55
massalis 34
megalospora 43
melampodii 25, 34-35, 37
menthae 297
mikanae 28
minussensis 28
mirifica 34
moliniicola 51
morigera 51
mundula 22
nanomitra 34
neocoronata 51
nesodes 60
noccae 34
nociva 44
nocticolor 43
notha 40
nothoscordi 21
notopterae 35
nuda 35
oahuensis 51
obeisiseptata 29
obesispora 21
oblata 35
oblongula 44
obvoluta 44
okudairai 48
opaca 35
orbicula 28
ordinata 35
Osoyoosenis 44
otopappicola 35
otoppi 35
oxypetali 23
pachyspora 29
paederotae 60
pallor 22
paradoxica 51
parkerae 46
paroselae 45
parthenices 35
parthenii 30, 35
pazschkie

var. ferruginea 58

var. huteri 58

var. oppositifolia 58
pedatissima 60
perfuncta 47
perincerta 24
permagna 62
permagnum 20
perscita 47
perspicabilis 24
pertrita 40
pervenusta 24
pestibilis 40
petalstemonis 77
picturata 49
pimpinellae

ssp. carlottae 22

a17

pinaropappi 28
pinguis 40
piqueriae 29
plagianthi 49
plumbaria 54
poae-nemoralis

ssp. hyparctica 51
poecilochromae 61
polemonii 54
polliniae-quadrinervis 51
polygoni-amphibii

var. polygoni-amphibii 297
polypogonis 291, 298
polymniae 35
poromera 22
porphyretica 47
porphyrogenita 43
praealta 40
praeculta 25
praegracilis

var. cabotiana 51

var. connersii 51

var. praegracilis 51
praemorsa 29
praetermissa 36
praevara 21
proba 36

' procerula 38

proluviosa 38
punctoidea 36
puritanica 44
rangiferina 52
rata 40
recondita 298
repentina 22
retecta 55
ribesii-japonica 46
rimosa 292
ripulae 25
rudbeckiae 36
ruderaria 25
rufescens

var. densiflora 61

var. rufescens 61
russa

var. leptotricha 25
sakamotoi 46
salebrata 25
sana 47
saxifragae

var. heucherarum 58

var. holochloe 58

var. longior 58

var. mitellae 58
saxifragae-gei 58
saxifragae-micranthae 57
schedonnardi 52
schileana 36
schistocarphae 38
scillae-rubrae 52
scimitriformis 48
scirpi 299

518

[Puccinia ] scleropogonis 52

semota 36
senecioicola 39
senecionis 39
seposita 48
serratulae 27
setariae 52
shiraiana 3, 21
silphii 36
solidpes 29
sonorae 36
sonorica

var. sonorica 52
sparganioides 52
sphacelicola 48
sphenica 25
splendens 36
stapfiolae 299
Steiractiniae 36
stephanomeria 27
striiformis 291

var. dactylidis 300

var. striiformis 299
subaquila 36
subglobosa 37
subita 38
substerilis

var. oryzopsidis 52

var. substerillis 52
substriata 52
suksdorfii 28
takakumensis 48
tanaceti 32
tecomicola 21
terrieriana 296
tetramerii 21
thelypodii 42
thuemenia 61
tithoniae 37
tosta luxurians 50
tottoriensis 58
tournefortiae 41
treleasiana 55
triumfettae 61
trixitis 38
troximontis 28
turgidipes 37
tuyutensis 300
unica 52
urbanis 55
vaga 37
vanillosmopsidis 40
velata 45
veniabilis 40
verbesinae 37
vergrandis 45
veronicae-longifoliae 61
veronicarum 60
vexans 53
vilfae

var. mexicana 53

var. vilfae 53

vinulla 62
violae

ssp. americana 62
virgata 53
wedeliae 37
wedeliicola 37
wulfeniae

ssp. saximontana 61
xanthit 37
ximenesiae 37
yosemitana 54
zaluzeniae 38
zexmeniae 35, 38

Pucciniastrum

fagi 63
hikosanense 63

hydrangeae-petiolaridis 63

kusanoi 63

miyabeanum 63

pseudo-cerasi 63

styracinum 64

yoshinagai 64
Pucciniosira

holwayi 64
Punctelia

subrudecta 426-427
Pycnoporellus 389
Pyrenochaeta

xanthoriae 425
Pyrofomes 305-306, 314

albomarginatus 306-307, 314

aurantiacus 306, 314

demidoffii 188, 306, 314

fulvo-umbrinus 314

lateritius 305-310, 313-314
perlevis 305-307, 311-314

tricolor 306-307, 314
var. eumorphus 307
var. indicus 307
var. shoreicola 307
var. tricolor 307
Pythium 156, 172
aphanidermatum 157
proliferum 156

ret)

Quadracaea 103, 106-107

mediterranea 103, 104-105, 107-109

Ramalina
fastigiata 422-423
yemensis 418
Ramocladia
reticulata 152
Ravenelia
annulata 64
bizonata 64
cumminsii 64
distans 64
echinata
var. ectypa 65
ectypa 65
expanse 65

[Ravenelia] faceta 65
idonea 65
irregularis 65
rata 65
sololensis 65
theiseniana 65

Remispora
crispa 398-400, 408
galerita 398, 400, 408
maritima 398, 408
pilleata 398, 408
quadriremis 398, 408
spinibarbata 398, 408
stellata 398, 409

Repetobasidiellum
fusisporum 185

Rhizidiomyces
bivellatus 156

Rhizidium
endosporangiatum 167
megastomum 167
microcystidis 167
parasiticum 167
renifore 168
tomiyamanum 168

Rhizoclosmatium
hyalinum 168

Rhizophlyctis 168
aurantiaca 168
boninensis 168
bonseyi 168
columellae 168
costatus 168
fuscus 163, 168
hirsutus 168
ingoldii 168
lovettii 168
mastigotrichis 168
oceanis 168

var. floridaensis 168
petersenii

var. appendiculata 168
reynoldsii 168
rosea 152, 154, 168
serpentina 168
spinosa 161
tropicalis 168
variabilis 168

var. burmaensis 168
willoughbyi 168

Rhizophydium 164, 168
achnanthis 160, 168
aestuarii 164, 168
anatropum 164, 168
androdioctes 169
angulosum 169
annulatum 169
aphanomycis 169
aureum 169
biporosum 169
blastocladianum 169
blyttiomycerum 169

brevipes

var. marinum 164, 169
bumilleriae 164, 169
capillaceum 169
clavatum 169
coleochaetes 154, 169
collapsum 169
conchiforme 169
condylosum 169
contractophilum 169
coralloidum 169
coronum 172
difficile 160, 169
digitatum 166, 169
dubium 169
echinocystoides 169
ellipsoidium 164, 169
elyensis 169
fragilariae 160, 170
fugax 170
gonapodyanum 170
hispidulosum 170
horizontale 170
karlingii 170
lagenaria 170
laterale 170
lenelangeae 170
littoreum 170
macroporosum 170
manoense 170
melosirae 160, 170
mougeotiae 170
nobile 170

novae-zelandiensis 164, 170

obpyriformis 164, 170
olla 164, 170
patellarium 170
pedicellatum 170
pelagicum 170
piriformis 160, 170
planktonicum 170, 173
poculiforme 170
polystomum 170
porosum 170
proliferum 170
punctatum 171
rarotonganensis 171
reflexum 171
rhizinum 171
rotundum 171
scenedesmi 164, 171
sibyllinum 171
signyense 171
skujai 164, 171
sparrowii 171
sphaerocarpum

var. rhizoclonii 171

var. spirogyrae 171
sphaerotheca 171
spinosum 149, 171
spinulosum 164, 171
squamosum 171

519

520

[Rhizophydium] stellatum 171
stipitatum 171
tenue 164, 171
tetragenum 171
tubulatum 171
ubiquetum 171
undatum 171
undulatum 171
urcelolatum 171
venezuelensis 172
venustum 172, 174

Rhizoplaca 423
chrysoleuca 423

Rhodocybe
mundula 220
truncata 220

Rhodophyllus
lanicus 216
sinuatus 216

Rigidoporus 387, 389

Rimelia
cetrata 418
reticulata 426-427

Roestelia
brucensis 65

Royoporus 143, 147
spathulatus 145, 147

Rozella 172
blastocladiae 172
coleochaetis 172
diplophlyctidis 172
itersoniliae 172
longicollis 172
longisporangia 172
parva 172

Rozellopsis 172

Ruminomyces 151, 172
elegans 151, 172

Russula 471-472, 475, 477-478

olivaceoviolacens 473

Sarocladium
oryzae 209
Savoryella
lignicola 398, 409
Schenella 82, 100-101
microspora 82
simplex 79, 81-82, 88, 92
Scherffeliomyces
appendiculatus 157, 172
leptorrhizus 157, 172
Scherffeliomycopsis 172
coleochaetis 172
Scirrhia
acicola 437
pini 437
Sclerococcum
parmeliae 415, 425-426
serusiauxii 424-425
simplex 427
Scolecobasidiella 260
tropicalis 260, 262

Scorpiosporium 455, 463
angulatum 464
anomalum 464
gracile 464
minutum 455, 463-464

Scutellospora 130, 135

cerradensis 129, 130-132, 134-136

coralloidea 135
fulgida 135
gilmorei 135
gregaria 135
heterogama 135
pellucida 135
persica 135
verrucosa 130, 135
Scytinostromella
heterogenea 186
Sebacina 118
epigaea 111, 118-119
Selenosporella 103-104, 106-107
Septochytrium
willoughbyi 172
Septosperma 172
anomala 172
anomalum 172
irregulare 151, 172
multiforma 172
multiforme 172
rhizophidii 172
rhizophydii 172
spinosa 172
spinosum 172
Serendipita
vermifera 111, 120
Sigmoidea
luteola 398-399, 410
Siphonaria
variabilis 172
Sistotrema
subtrigonospermum 186
Skierka
holwayi 65
Sorateae
holwayii 66
Sorochytrium 172
milnesiophthora 173
Sparrowia 173
parasitica 173
subcruciformis 173
Sparrowmyces 173
sparrowil 173
Sphaerella
myricariae 439
Sphaeria
myricariae 439
Sphaerita
dinobryi 173
Sphaeromonas
communis 153
Sphaerophragmium
everinum 66

Sphaerulina
oraemaris 398-399, 409
Spiculogloea 111-112
occulta 111-113, 114
Spizellomyces 173
acuminatus 164, 173
dolichospermus 173
kniepii 165, 173
lactosolyticus 173
palustris 165, 173
plurigibbosus 166, 173
pseudodichotomus 173
punctatus 165-166, 173
Sporidesmium 278
Sporophlyctidium
neustonicum 173
Sporothrix 209
vizei 209
Stemonitis 91, 100
axifera 85-86
var. axifera 84-85
var. smithii 85-86
confluens 86-88
flavogenita 93-94
foliicola 91
fusca 87
herbatica 91
hyperopta 79, 89-90, 92, 99
microsperma 79, 90
nigrescens 90-92, 99
pallida 91, 99
splendens 93-94
virginiensis 90
Stemonitopsis
hyperopta 89
subcaespitosa 98, 100
Stigmina 435
concentrica 435
Strigula 323, 326
affinis 326
australiensis 326
johnsonii 326
minutula 326
occulta 323-326
stigmatella
var. alpestris 326
taylorii 323, 325
Strobilurus
stephanocystis 220
Stropharia
aeruginosa 218
coronilla 218
semiglobata 218
stercoraria 218
Subulispora
procurvata 373, 381-382
rectilineata 373, 382-384
Suillus
bellini 215
boudieri 215
bovinus 215
granulatus 215

luteus 215

pictus 215

variegatus 215
Symphytocarpus

confluens 86-87
Synchytrium 149-150, 161, 173

grandis 161

namae 163

Talaromyces 476
Thekopsora
pseudo-cerasi 63
Thermoascus 225-226, 228, 230
aegyptiacus 230, 232, 238
aurantiacus 226, 230, 232-235
var. levisporus 230
crustaceus 226, 230, 232, 235-239
taitungiacus 225-226, 229-232
thermophilus 230, 232
Thielavia 241, 244
aegyptica 244
appendiculata 244
arenaria 244
australiensis 244
emodensis 244
expansa 244
fragilis 244
heterothallica 244
hyalocarpa 244
hyrcaniae 244
microspora 244
minuta 244
octospora 244
peruviana 244
pingtungia 241, 242-246
subthermophila 244
terrestris 244
tetraspora 244
tortuosa 244
Thozetella
cristata 373, 382-384
Ticogloea 260
guttulata 260, 262
Tofispora
repetospora 181
Torpedospora
ambispinosa 398-399, 409
Torrendia
pulchella 217
Trachypus
pseudoscaber 215
Trametes 320
feei 320
suaveolens 320
Tremella 112
foliacea 123
mesenterica 123
Tricella
acuminata 19
Trichocladium
achrasporum 398, 410
alopallonellum 398-399 ,410

a2

22

[Trichocladium] constrictum 398-399, 411

Tricholoma
albobrunneum 220
aurantium 220
auratum 220
batschii 220
caligatum 220
flavovirens 220
grammopodium 220
melaleucum 220
orirubens 220
populinum 220
rutilans 220
sejunctum 220
sordidum 220
terreum 220
virgatum 220

Tricladium 455, 463
attenuatum 460
minutum 455, 464, 468

Triparticalcar 173
arcticum 164, 173

Triphragmiopsis
isopryri 66

Triphragmium
isopryri 66
koelreuteriae 17

Triposporium 275, 278
deviatum 278
elegans 278

Tritirachium
oryzae 288

Tubaria
furfuracea 220
pellucida 220
trigonophylla 220

Tuber 263, 365, 370-371
candidum 365, 368
excavatum 370
lyonii 365-367, 369-371
malacodermum 371
melanosporum 371
rufum 365, 368, 370-371

var. nitidum 368, 370-371

var. rufum 371
spinoreticulatum 371
texense 365-366, 368, 370

Tulasnella 122
balearica 111, 121-122
brinkmannii 122
calospora 122
deliquescens 122
pallida 123
pallidocremea 122
violea 123

Tylopilus
felleus 215

Uredinopsis
americana 66
arthurii 66
coplandii 66

kameiana 66
longimucronata 67
struthiopteridis 67
woodsiae 67
Uredo

adianti-capilli-vernis 14
apecheca 68
aztechana 68
cancellata 292
cenchricola 293
cyperi-tegetiformis 294
detecta 67
eleusine-indicae 293
glumarum 299
imperatae 295
laeviuscula 17
linearis

var. b polypodi 14
lini 15
nassauviae 67
nootkatense 13
operculina 67
phormi 68
polypogonis 298
schizachyrii 294
scopelandii 16
sphacelicola 68
spirostachydis 67
velata 45
zeugitis 68

Uromyces 3, 291

actinostemonis 71
americanus

var. californicus 70
amoenus 69
amurensis 72
anguriae 70
armeriae

ssp. hudsonica 74

ssp. pacificus 74
asperulae 76
aspiliae 69
azukicola 72
bauhiniicola 72
beckmanniae 74
bisbyi 75
bradburyae 72
cisneroanus 71
clitoriae 72
claytoniae 75
cologaniae 72
croci 73
digitatus 74
dolichosporus 69
galii 76
halstedii 74
hardenbergiae 72
hedysari-paniculati 72
inaequialtus 70
intricatus

var. major 75

var. nivea 75

[Uromyces intricatus] var. pluriporosus 75
var. umbellati 75
lapponicus
var. oxytropis 72
lezpedezae-procumbentis 72
limonii-caroliniani 74
linearis 300
lineolataus
ssp. nearcticus 71
loranthi 74
manihotus-catingae 71
mexicanus 72
neurocarapi 73
nothoscordi 68
novissimus 70
oaxacanus 71
oblectaneus 71
occidentalis 73
panici-sanguinalis 75
peckianus 75
ssp. rugulosa 75
peracarpae 69
pereskiae 73
polycnemi 70
primaverilis
ssp. nothoscordi 68
ssp. primaverilis 69
probus 73
psychotriae 76
ratus 70
ribicola 73
rostratus 73
rumicis 301
salpichroae 76
scutellatus 71
senecionicola 69
setariae-italicae 301
shikokianus 73
sisyrhynchii 73
socius 74
sparganii
ssp. asiaticus 77
sphaericus 69
tolerandus 71
tranzschelii 71
trifolii-repentis 302
var. trifolii-repentis 301
unitus 76
ssp. montanensis 76
ssp. pacificus 76
ssp. spraguae 76
ssp. unitus 76
vicinus 70
Uromycladium
bisporum 77
simplex 77
Uropyxis
arisanensis 77
diphysae 77
holwayi 77
petalstemonis 77
wootoniana 11

0 9 a6

Usnea 422-423
florida 418

Varicosporium
elodeae 463
Verticillium
luteo-album 209
Vesiculomyces
epitheloides 182
Volvariella
bombycina
var. maxima 216
media 216
speciosa 216
Vuilleminia 181, 187
comedens 186-187
cystidiata 187
macrospora 187
megalospora 187
obducens 181, 186-187
subglobispora 187

Wettsteinina 434
yuccigena 434

Xanthoria 421
candelaria 421
parietina 421-422, 425
Xenasma
pruinosum 188
pulverulentum 188
rimicolum 188
Xenostele
takakumensis 48
Xerocomus
badius 215
subtomentosus 215
Xeromphalina
campanella 220
Xylaria 327, 338, 352, 356
allantoidea 327-330
apiculata 352
arbuscula 352-354, 356-358
badia 327, 329-330, 332
castorea 344
corniformis 337
cubensis 327, 330, 332-334, 336, 338,
342
curta 336-337, 354
feejeensis 327, 337, 339, 344, 354
frustulosa 338, 341
grammica 327, 339-340, 342, 346, 348,
35057352
laevis 327, 334, 336, 342
luteostroma 344
luteostromata 327, 343-344
mellisii 358
multiplex 327, 344, 346-348, 350, 352
pallida 327, 346, 348, 352
papulis 327, 347-348, 350, 352
poitet-<327, 330,333, 352
polymorpha 356

524

[Xylaria ] schreuderiana 352
schweinitzii 351, 354, 356
Xylocoremium
flabelliforme 336

Zalerion

maritimum 398, 411

varium 398, 411
Zeloasperisporium 284-285

hyphopodioides 283-285, 286-287
Zetiasplozna

cordylines 199

Zygogloea 112
Zy gorhizidium

affluens 172-173
asterionellae 173
chlorophycidis 173
cystogenum 174
melosirae 174
planktonicum

f. asterionellae 174

f. synedrae 174
vaucheriae 174
venustum 172, 174

Reviewers, Volume Sixty

The Editors express their appreciation to the following individuals who have, prior to
acceptance for publication, reviewed one or more of the papers appearing in this volume:

T. J. Baroni G. L. Hennebert
D. J. S. Barr Y. Hiratsuka

M. E. Barr E. Horak

G. H. Boerema Y.-M. Ju

K. L. Braun H. W. Keller
H. H. Burdsall, Jr. B. Kendrick

R. F. Castafieda Ruiz J. W. Kimbrough
G. B. Cummins J. J. Kohlmeyer
E. Descals C. Lado

W. Gams T. Less@e

E. B. Gareth Jones M. J. Larsen

R. L. Gilbertson D. Malloch

J. H. Ginns D. W. Mitchell
J. Guarro J. B. Morton

R. E. Halling D.N. Pegler

D. M. Henderson D. H. Pfister

M. Rajchenberg
A. W. Ramaley
P. Roberts

J. D. Rogers
A. Y. Rossman
C. Roux

K. A. Seifert

. Sérusiaux

. A. Shearer

. V. Subramanian
. Triebel

. E. Tulloss

. Walker

. J. K. Wang
_ Webster

. E. Wright

“=QAQAFTNIAM,

Publication Date Volume Fifty-Nine

MYCOTAXON _ for July-September 1996, volume 59: 1-524
was issued on September 18, 1996.

MYCOTAXON T-SHIRTS NOW AVAILABLE

MYCOTAXON

the journal of fungal systematics front

bad taxonomy

&} i
can
KILL

You may contact Mycotaxon, Ltd.
by e-mail at <rkorf@innet.com>, or by fax at (607) 273-4357,
or by mail at Mycotaxon, Ltd., P.O. Box 264. Ithaca, NY 14851
for prices and ordering information

Sizes: M, L, and XL, for chest sizes 38-40, 42-44, and 46-48 inches,
in 100% pre-shrunk cotton, maroon lettering on white shirt

Reduced price for Mycotaxon subscribers

EDITORS of MYCOTAXON

JEAN BOISE CARGILL, Editor-in-Chief
Harvard University Herbaria, 22 Divinity Ave.
Cambridge, MA 02138, U.S.A.

ASSOCIATE EDITORS

Karen Gettelman David S. Hibbett G. L. Hennebert Pavel Lizon
Index Editor Book Review Editor French Language Editor Asst. to the Ed.in-Chief
Indiana University Harvard Univ. Herbaria, 32 Rue de l'Elevage Plant Pathology Herbarium,
Bloomington, IN Cambridge, MA USA B-1340 Ottignies - LLN Corel! Univ.
Belgium Ithaca, NY USA

OB ale eg age a neo SD a as UR Ne nen LEITGRS TIPCRESART Ce ROE
MYCOTAXON. is a quarterly journal devoted to all phases of mycological taxonomy and nomenclature. All
articles are reviewed by specialists prior to acceptance. Publication is open to all persons. Papers may be in
French or in English. Summaries in those or any additional languages desired by the authors are given for
longer articles. Printing 1s on high quality, acid-free book paper. Authors prepare their own camera-ready
copy. after having received critical comments from pre-submission reviewers. Instructions to Authors appeared
‘n MYCOTAXON 50: 493-497, 1994. A copy of these instructions will be sent upon request to the Editor-in-
Chief.

SUBSCRIPTION INFORMATION
Each volume, beginning with volume 3, contains at least 512 pages, and consists of an irregular number of
quarterly issues (rarely an additional issue, a Festschrift, may be included in a volume). Each issue of
MYCOTAXON varies in nuimiber of pages. Subscriptions are normally on a per volume basis, but subscribers
may choose an annual basis to avoid frequent billing. Currently this would involve prepaying three volumes.
Personal subscriptions ave available at a substantially reduced rate for individuals who agree not to deposit
(heir copies in another library than their personal one within 3 years of receipt. Address all orders to the
Mycotaxon Order Department, not to the Editors: Prices beginning with volume 53 are:

USA Canada/Mexico Other Foreign (Air)
REGULAR (multiuser) $65.00 $67.00 US $70.00 US
PERSONAL (individual) $30.00: $32.00 US $37.00 US
(All back volumes are still available. All complete volumes ate available at $30.00 per volume when
shipped by surface mail, $45 per volume by air mail, except vols. 1, 34 and 46 available only in

complete runs.)

Place subscriptions through the ORDER DEPT., MYCOTAXON, LTD., P. O. BOX 264, ITHACA, NY 14851-
0264, U.S. A. or through your agent. MYCOTAXON may also be obtained on a journal-exchange basis. This
may be arranged with journals, institutions, or individuals who have difficulty in obtaining foreign currency.

TWENTY-VOLUME CTIMULATIVE INDICES, 1974-1984 & 1984-1991
MYCOTAXON CUMULATIVE INDEX FOR VOLUMES LXX (1974-1984) by Richard P. Korf & Susan C.
Gruff (ISBN 0-930845-00-5) is available at $17.50 postpaid, and MYCOTAXON CUMULATIVE INDEX FOR
VOLUMES XXI-XL (1984-1991) by Richard P. Korf & Susan C. Gruff (ISBN 0-930845-01-3) is available at
$30.00 postpaid, MY COTAXON, LID., P.O: BOX 264, ITHACA, NY, 14851-0264, U.S.A.

AVAILABILITY IN MICROFORM, TEAR SHEET & PHOTOCOPY
MY COTAXON is also available in microfiche and in microfilm from University Microfilms, 300 North Zeeb
Road. Ann Arbor, MI 48106, U.S.A., or 30-32 Mortumer Street, London WIN 7RA, England, from whom prices
may be obtained. Tear sheets or photocopies of individual articles may be obtained through The Genuine
Article!M 1.8 1.,3501 Market Street, Philadelphia, PA 19104, U.S.A:

RECENT BOOK PUBLISHED BY MYCOTAXON, LTD.
FUNGI OF CHINA by S. C. Teng (1996), edited by Richard P. Korf, 586+xiv pp., cloth-bound, 8-1/2 x 11
isches, 426 ilhustrations, 1 map, 1 photograph, $79 postpaid.

CONTACTING MYCOTAXON'S ORDER DEPARTMENT BY E-MAIL OR FAX
To reach the Order Department for information or placing orders, you may E-Mail to rkorf@innet.com or you
inay FAX to Richard Korf at (607) 273-4357.

NEW! VISIT MYCOTAXON ON WORLD WIDE WEB AT www-mycotaxon.com

wy

s